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'dens’ Bulletin Singapore Vol. 67(1) 2015

Singapore

ISSN 0574-7859

Singapor^'^otanic Gardens

THE GARDENS’ BULLETIN, SINGAPORE

The Gardens^ Bulletin, Singapore is a peer-reviewed journal publishing original papers and
reviews on a wide range of subjects: plant taxonomy (including revisions), phytogeography,
floristics, morphology, anatomy, as well as horticulture and related fields, such as ecology and
conservation, with emphasis on the plant life of the Southeast Asian-Pacific region.

Dr David J. Middleton Dr Jana Leong-Skomickova
(Editor-in-Chief) (Managing Editor)

S. Lee Y.W. Low Christina Soh

(Graphics Editor) (Copy Editor) (Business Manager)

Editorial Advisory Board

Professor Sir Peter Crane
Yale University
USA.

Dr Rogier P.J. de Kok
Royal Botanic Gardens, Kew
UK.

Dr W. John Kress

National Museum of Natwal History
Smithsonian Institution
U.S.A.

Dr Mark Hughes

Royal Botanic Garden Edinburgh
UK.

Dr Kiat W. Tan
Gardens By The Bay
Singapore

Dr Nigel P. Taylor
Singapore Botanic Gardens
National Parks Board, Singapore

Dr Ian M. Turner
Royal Botanic Gardens Kew
U.K

Dr Jan-Frits Veldkamp
National Biodiversity Center
The Netherlands

Dr Jun Wen

National Museum of Natural History
Smithsonian Institution
U.S.A.

Professor Nianhe Xia
South China Institute of Botany
PR. China

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Instructions for contributing authors (continued)

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Herbarium abbreviations follow

Thiers, B. (2013). [continously updated] Index Herbariorum: A global directory of public herbaria and
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Journal and book title abbreviations. For journals:

Bridson, G.D.R., Townsend, S.T., Polen, E.A. & Smith, E.R. (eds) (2004). BPH-2: periodicals with
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Institute for Botanical Documentation.

For books:

Stafleu, F.A. & Cowan, R.S. (eds) (1976-88). Taxonomic Literature. 2nd edition. 7 vols [Regnum Veg.
vols 94, 98, 105, 110, 112, 115, 116].

Stafleu, F.A. & Mennega, E.A. (eds) (1992-). Taxonomic Literature. Supplements [Regnum Veg. vols
125,130,132].

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References listed at the end. There, works mentioned in the text are listed alphabetically as follows:
Dallwitz, M.J., Paine, T.A. & Zurcher, E.J, (1999). User’s Guide to the DEETA Editor, http://
biodiversity.uno.edu/delta/ (accessed on 2 Aug. 2010).

Persson, C. (2000). Phylogeny of Gardenieae (Rubiaceae) based on chloroplast DNA sequences from the
rps 16 nitron and P‘«L(UAA)-F(GAA) intergenic spacer. Nordic J. Bot. 20: 257-269.

Ridley, H.N. (1930). The Dispersal of Plants Throughout the World. Ashford, U.K.: L. Reeve.

Smith, A.C. & Daiwin, S.P. (1988). Rubiaceae. In: Smith, A.C. (ed) Flora Viliensis Nova, A New Flora
q/T/// 4: 143-193.

References to web-based resources should include either a doi (digital object identifier) specification
or fiill URL mentioning also the date it was accessed. Use of DNA sequences from GenBank should be
acknowledged and the studies for which the sequences were generated should be cited.

Style of nomenclatural summaries. The following style is required:

Ornithoboea arachnoidea (Diels) Craib, Notes Roy. Bot. Gard. Edinburgh 11: 251 (1920); Burtt, Notes
Roy, Bot. Gard. Edinburgh 22: 294 (1958).

Ornithoboea parishii C.B.Clarke in A.DC. & C.DC., Monogr. Phan. 5(1): 148 (1883).

If authors include full bibliographic data for these works in the list of references at the end of the paper,
they should also be mentioned in the text briefly, e.g., “Nomenclatural references researched include
Blume (1849) and Ro-xburgh (1 824).”

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Front cover picture: Hanguana rubinea (Photo by Jana Leong-Skomickova)

The Gardens’ Bulletin

Singapore

VOL. 67(1) 2015 ISSN 0374-7859

CONTENTS

J. Leong-§kornickova & P.C. Boyce

Hanguana in Singapore demystified: an overview

with descriptions of three new species and a new record 1

M. Rodda, D. Lee & S. Mishak

Dischidia acutifolia (Apocynaceae, Asclepiadoideae)

a new record for the Singapore flora 29

I.M. Turner

Yvom Ancistwcladus to Tristaniopsis via Tetramerista

- the convoluted history of a Wallich collection and its impact

on the native flora of Singapore 35

S. Lindsay & D.J. Middleton

The clarification and synonymisation of two taxa of Vittaria
from Peninsular Malaysia and a new combination

in Haplopteris (Pteridaceae subfam. Vittarioideae) 39

A.R. Rafidah, S. Syahida-Emiza, R. Kiew & K. Imin

Bothriospermum (Boraginaceae), a new generic record

for the Flora of Peninsular Malaysia 45

A. A. Joffre, A.K. Muhammad Ariffin, Y.W. Low & K.M. Wong

Novitates Bmneienses, 1 . A background to the botanical survey of Brunei
Darussalam, and a new species of Jarandersonia (Malvaceae) 51

Y.W. Low, A.A. Joffre & A.K. Muhammad Ariffin

Novitates Bmneienses, 2. A remarkable new species oi Begonia
sect. Petermannia (Begoniaceae) from Bmnei Damssalam

61

K. M. Wong & Y.W. Low

Novitates Bruneienses, 3. Eight new woody plants

in the Brunei flora, including five new species 69

S. Rahayu, U. Meve & M. Rodda

Hoya undulata (Apocynaceae, Asclepiadoideae), a new myrmecophytic

species from Borneo, and typification of H. darwinii 85

N.K.E. Undaharta, I Made Ardaka, Agung Kurniawan & Bayu Adjie

Begon ia bimaensis, a new species of Begonia

from Sumbawa Island, Indonesia 95

C. Puglisi, S. Suddee, P. Triboun & D.J. Middleton

Anew species of Paraboea (Gesneriaceae) from Thailand 101

H. Kurzweil & Saw Lwin

New orchid records for Myanmar, including

the first record of the genus Stereosandra 107

M.F. Newman

A new species of Zingiber (Zingiberaceae) from Lao P.D.R 123

L. Bai, J. Leong-Skornickova & N.H. Xia

Taxonomic studies on Zingiber (Zingiberaceae) in China I: Zingiber kerrii

and the synonymy of Z. menghaiense and Z. stipitatum 129

C. Deori, N. Odyuo & A.A. Mao

Pennilabium labanyaeanum (Orchidaceae),

a new species from Meghalaya, Northeast India 143

K. Thoiba, A.K. Pradeep & C.N. Sunil

Tripogon bimucronatus (Poaceae: Chloridoideae: Tripogoninae),
a new species from India I5I

K. Bransgrove & D.J. Middleton

A revision of Epithema (Gesneriaceae) 159

B.R. Maslin

Synoptic overview of Acacia sensu lato (Leguminosae: Mimosoideae)
in East and Southeast Asia

231

BOOK REVIEW

Flowering Plants of the Western Ghats, Volume 1 (Dicots)

T.S. Nayar, A. Rasiya Beegam & M. Sibi. 2014 & Flowering Plants
of the Western Ghats, Volume 2 (Monocots). T.S. Nayar, M. Sibi
& A. Rasiya Beegam. 2014. (J. Leong-Skomickova)

Date of publication: 4 June 2015

Copyright ©

National Parks Board
Singapore Botanic Gardens
1 Cluny Road
Singapore 259569

251

Printed by Oxford Graphic Printers Pte Ltd

Gardens’ Bulletin Singapore 67(1): 1-28. 2015
doi: 10.3850/S2382581215000010

1

Hanguana in Singapore demystified: an overview with
descriptions of three new species and a new record

J. Leong-Skomickova^ & P.C. Boyce^

^Herbarium, Singapore Botanic Gardens, National Parks Board,

1 Cluny Road, 259569, Singapore
jana_skomickova@nparks.gov.sg

^Institute of Biodiversity and Environmental Conservation (IBEC),
Universiti Malaysia Sarawak, 94300 Samarahan, Sarawak, Malaysia

phymatarum@gmail.com

ABSTRACT. The genus Hanguana (Hanguanaceae - Commelinales) was until recently
believed to be represented in Singapore by a single species, Hanguana malayana. Recent
extensive surveys, coupled with the detailed study of fresh and herbarium material, however,
suggests the presence of six species. In addition to the recently described Hanguana neglecta,
three additional species, H. corneri, H. rubinea and H. triangulata, are described here as new to
science. Of the tliree newly described species, Hanguana rubinea and//, triangulata are native,
while H. corneri is believed to have been introduced from Peninsular Malaysia to Singapore
and planted in Bukit Timah Nature Reserve in the 1930s. Hanguana nitens is recorded for
the first time fi*om Singapore and is also certainly native. Wliile a large helophytic species of
Hanguana, currently interpreted to be Hanguana malayana, is cultivated as an ornamental
plant in Singapore, no native populations have been observed by us in the wild and neither do
any herbarium records exist to suggest that this species was ever native to Singapore. Colour
plates and a key to all Hanguana species are provided. Notes on habitats and preliminary lUCN
assessments are also included for all native species to better facilitate conservation efforts of
Hanguanaceae in Singapore. The existence and taxonomic potential of scale structures sheathing
the inner staminodes in female flowers, named here as staminodial scales, are highlighted here
for the first time. The nnportance of seed characters is also discussed.

Keywords. Bukit Timah Nature Reserve, Central Catchment Nature Reserve, Hanguana
corneri, H. malayana, H. neglecta, H. nitens, H. rubinea, H. triangulata, seeds, staminodial
scales

Introduction

Hanguanaceae (Commelinales) is a small monogeneric family distributed in S and
SE Asia, with the highest diversity in the Sunda region. The family is perhaps the
most understudied family of monocots (Siti Nurfazilah et al, 2011), deserving much
closer attention in the region. Currently 1 1 species are recognised of which eight were
described in the last five years (Siti Nurfazilah et al., 2010, 201 1; Mohd Fahini et al.,
2012; Niissalo et al., 2014). We, however, estimate that the number of species is likely
to exceed 50 as future exploration and detailed studies, particularly of mature fruiting
material, progress.

2

Gard. Bull. Singapore 67(1) 2015

While Singapore has the best-known flora in SE Asia owing to the small size of
the country, coupled with long-tenn botanical exploration (Niissalo et ah, 2014), the
state of Icnowledge is still far from complete. Numerous new records (e.g,, Low et ah,
2014; Rodda et ah, 2012), as well as descriptions of new taxa (Sugumaran & Wong,
2014; Leong-Skomickova et ah, 2014), including Hanguana neglecta (Niissalo et ah,
2014), have recently been published from Singapore.

All existing herbarium records of Hanguana at SfNG, collected in Singapore
between 1885 and 2009 (but excluding our recent collections) and amounting to
20 specimens in all, were previously misidentified as Hanguana malayana (Jack.)
Men*., a name much misapplied across Asia. As such, Hanguana malayana is the only
species listed in works dealing with the Singapore flora (Keng et ah, 1998; Chong et
ah, 2009). During extensive fieldwork since 201 0 by the first author, it was observed
that several Hanguana species are present in Singapore's forests rather than just the
single species previously suggested. Further fieldwork, coupled with the expertise of
the second author who has been working on the Hanguanaceae of the Sunda region
since 2010, revealed that at least six species are present in Singapore, although only
four are considered native. These include Hanguana neglecta recently described from
Singapore (Niissalo et ah, 2014), H. nitens a species recently described from Johor,
Peninsular Malaysia (Siti Nurfazilah et ah, 2010), and two new species described
below as H. mbinea and H. triangulata. There is no confirmed record of Hanguana
malayana ifom the wild, be it by direct observation, or through the existence of even
a single herbarium specimen from Singapore. Hanguana corneri, while described
here as new to science, has been introduced to Bukit Timah Nature Reserve, and is not
considered a Singaporean native.

The difficulty of studying species of Hanguana from herbarium material alone,
and the importance of infmctescence structure, stigma, fruits, seeds and indumentum
for delimitation of taxa, has been highlighted before by Siti Nurfazilah et ah (2010,
2011). The importance of the morphology of the stigma cannot be overstated as it
preserves well in herbarium material. The position of the stigma in relation to
fruit, however, still requires further study. We have observed that in some species
(e.g., Hanguana mbinea and H. triangulata^ both described below), the position
of the stigma on ripe fruit (oblique or central) is influenced by the number of seeds
developing in the berry. Hanguana berries are trilocular, with a single ovule in each
locule, of which one, two or all three can develop. The actual number of seeds which
eventually develops presumably depends on pollen availability, but also on species
strategy. While the number of seeds which develop per berry is more or less stable in
some species (e.g., always single mH. neglecta), it can vary in others (e.g., one or two
in H. nitens, H. mbinea and H. triangulata', one, two or thi'ee in H. malayana). While
seeds were previously acknowledged as a potential source of characters for Hanguana
taxonomy, the topic has not been discussed in detail. Descriptions or illustrations of
seeds are imfoitimately not available for many of the recently described species. Our
current study shows that seeds are extremely informative and this topic is, therefore,
discussed below. We also highlight the existence of previously overlooked structures
occurring in female flowers.

Hanguana in Singapore

3

The taxonomic importance of staminodial scales and seeds

Tillich (1996), Tillich & Sill (1999) and Rudall et al. (1999) made major contributions
to our understanding of the floral and seed structure of Hanguana. This then served as
a basis for recent descriptions and was further improved with the recent work of Siti
Nurfazilah et al. (2010, 2011) and Mohd Falimi et al. (2012).

Hanguana flowers are rather unifonn across the entire genus and seem to offer
few characters helpful in species delimitation. Male flowering material is rarely found
in the field (and, therefore, is also scarce in herbaria). The male flowers/inflorescences
are short-lived in comparison to female flowers, which progress to fruiting stage (a
process lasting several months) while retaining all floral parts practically unchanged or
slightly enlarged (e.g., tepals, staminodes, stigma). As fruits offer additional characters,
particularly the colour of the ripe fruit and seed morphology, Hanguana species are
most often described from fmiting material, a trend certain to be continued.

While examining fresh fruiting material of various species from Singapore,
the presence of scale-like stmctures sheathing the base of the three inner staminodes
was noticed. None of the previous publications dealing with Hanguana mentions
their existence (e.g. Airy Shaw, 1980; Tillich, 1996; Bayer et al., 1998; Tillich & Sill,
1999; Siti Nurfazilah et al., 2010, 2011), with the exception of Rudall et al. (1999)
who noticed them but misinterpreted them for inner staminodes. Further detailed
examinations of fresh as well as herbarium material, however, confirmed that six
staminodes are present in each flower (three outer and three umer) and that the three
scales are positioned at the base of the three imier staminodes, sheathing them partially
of fiilly. These scales differ in shape and in the presence of a hyaline margin (Fig. 1,
3B, 5B, 7B, 9B, 12B). They preserve well in herbarium specimens and are, therefore,
of good taxonomic potential, at least in certain species. While the exact origin and
function of these scales is not yet clear and requires further study, it has been suggested
(H.-J. Tillich, pers. comm.) that these might be a basal outgrowth of the staminodes,
possibly exuding a mucilage or a particular scent to attract a pollinator. We propose
here to use the term stammodial scales for these structures. Although staminodial
scales have been observed by us in all species we have so far examined from living
and herbarium material, it is not yet clear if these are indeed present in all Hanguana
species.

The seeds of Hanguana are unique in the monocots (H.-J. Tillich, pers. comm.).
They are more or less bowl-shaped with the hilum positioned at the base of the bowl,
the cavity being filled up by a placental tissue. Although the importance of seed size, the
varying depth of cavity and the degree of incurving of the rim were briefly mentioned
by Siti Nurfazilah et al. (2011), descriptions of seeds were unfortunately not provided
by Siti Nurfazilah et al. (2010) for most of the recently described species {Hanguana
exultans, H. pantinensis, H podzol icola, H stenopoda) or the large helophytic species
currently interpreted as Hanguana malayana.

From our recent work it is obvious that the morphological range of the seed
shape, and therefore their taxonomic potential, is considerable. The recently described
Hanguana neglecta (Niissalo et al., 2014) exhibits seeds which are almost round to

4

Gard. Bull. Singapore 67(1) 2015

f

Fig. 1. Detail of staminodial scales at the base of inner staminodes. A. Hanguana triangulata
Skomick. & P.C.Boyce. Note the presence of a hyaline margin on the staminodial scale. B.
Hanguana malayana (Jack.) Merr. Note the staminodial scale has no hyaline margin and is
composed of lobes. (Photos: Jana Leong-Skomickova)

Hanguana in Singapore

5

ovoid with a wedge-shaped ostiole. Similar seeds are also observed in Hanguana corneri
which is described below. The seeds of Hanguana rubinea and H. triangulata, also
described below, although both bowl-shaped, exhibit differently shaped appendages
along the proximal part of the rim. As previously mentioned, Hanguana species are
mostly collected in the fruiting stage, and fruits are often present in ample numbers
on the herbarium collections. Seeds, including small structures like appendages,
preserve well in dried material, as long as the fruits are collected fully mature or close
to maturity. It is possible to study even old material, from which the seeds can be
carefully extricated by removing the layer of fruit pulp after the berries are soaked
overnight in water at room temperature. In this study, seeds from specimens as old
as 1 889 collected by H.N. Ridley, were successfully extracted from fruits, providing
valuable confimiation of the determinations of Hanguana triangula ta and H. neglecta.

Hanguana in Singapore

The following account presents an identification key and colour plates to all six Han-
guana species so far known to be present in Singapore, including introduced and
cultivated species. Full descriptions are provided for the new taxa, while recently
described species are provided with notes to address any additions or deviations from
original or recent descriptions, these being clearly referenced. Based on sterile living
material we have observed in the field, we suspect the existence of at least one addi-
tional species, although field studies are needed to resolve its identity.

The tenninology used in the descriptions follows Beentje (2012) and the most
recent Hanguana works cited above. The type material of the species described from
ripe fruiting material also includes spirit material of fruits as well as extracted seeds.

Preliminary lUCN conservation assessments followed Davison et al. (2008) and
lUCN guidelines (lUCN, 2012) for local and global assessments respectively. Geo-
CAT (Bachman et al. 2011) was used to calculate EOO.

The native species are treated first in this account, followed by introduced and
cultivated species.

Key to Hanguana in Singapore

la. Large stoloniferous colonial herbs 2

lb. Solitary or clumping herbs lacking stolons 3

2a. Leaves stiffly erect with acute apex; lamina more or less flat or weakly irregularly
corrugate, semi-matt green; staminodial scales composed of lobes without hyaline
margin; stigma lobes large, flat, coimate at base, forming a bluntly triangular to

clover-leaf shape, almost obscuring the apex of ovary

H. malayana (sensu Siti Nurfazilah et al. 2010)

6

Card. Bull. Singapore 67(1) 2015

2b. Leaves weakly arching, with long-attenuate apex; lamina prominently corrugate,
shiny green with visible pattern of lighter and darker green (best observed on
young and medium aged leaves); staminodial scales entire with hyaline margin;

stigma lobes small, erect, separate, tear-shaped to obovate

H. nitens

3 a. Leaves green on both sides 4

3b. Leaves dark emerald-green above and dark red-purple underneath ....//. corneri

4a. Large herbs over 1 m in height; leaves arching; ripe fruits cream-white or ruby-

red; seeds bowl-shaped, more or less hemispherical 5

4b. Medium sized herbs not exceeding 0. 8 m in height; leaves spreading (not arching);

ripe fruits black; seeds y4 globose to ovoid with wedge-shaped opening

H. neglecta

5a. Lamina almost flat, abaxially with evenly distributed silky indumentum; ripe
fruits ruby-red; stigma lobes connate basally (sometimes imperfectly), with round

apices, foiming bluntly triangular structure; seed appendage acute

H, nibinea

5b. Lamina more or less conxigate, abaxially with unevenly distributed flocculose
indumentum; ripe fruits cream-white; stigma lobes with shaiply acute apices,
connate basally (always perfectly), fomiing sharply triangular structure; seed
appendage bluntly bilobed H. triangulata

Species native to Singapore

Hanguana neglecta Skomick. & Niissalo, Phytotaxa 188(1): 15 (2014). - TYPE:
Singapore, Bukit Timah Nature Reserve, slopes on lower end of Taban Loop along the
stream, 28 May 2014, Leong-Skornickovd, J. & Thame, A, JLS-2793 (holotype SING
(including spirit material); isotypes E, KEP, K). (Fig. 2, 3)

Ecology and distribution. In Singapore, Hanguana neglecta occurs on slopes in the
lowland primary tropical forest of Bukit Timah Nature Reserve, with increased density
closer to the stream. More than a century old herbarium records show that this species
also occun'ed in the Nee Soon and MacRitchie areas. Since it was described, we have
been able to re-locate a small population in the MacRitchie area {JLS-3099, deposited
at SING), but there are still no recent sightings of this species from the Nee Soon area.
The three currently known records from Peninsular Malaysia are confined to Johor
state, and are 56, 107 and 120 years old respectively.

Provisional lUCN conservation assessment. Following the criteria for national and
global conservation assessments (Davison et ah, 2008; lUCN, 2012), Hanguana

Hanguana in Singapore

7

Fig. 2. Hanguana neglecta Skomick. & Niissalo. A. Habit. B. Base of the plant showing semi-
aseending leafless stem (inset: detail of fruits, note the stigma composed of three obovate lobes
connate at base). C. Infructescence with fruits in various stages of ripeness (immature green to
glossy black mature fruits). From type JLS-2793. (Photos: Jana Leong-Skornickova)

8

Gard. Bull. Singapore 67(1) 2015

Fig. 3. Hanguana neglecta Skornick. & Niissalo. A. Detail of tepals (side view). B. Detail
of inner tepals, staminodes and staminodial scales. C. Detail of stigma on young fruit. D.
Longitudinal section of fruit, which is always single-seeded. E. Seed (top view). F. Seed (side
view). From type JLS-2793. (Photos: Jana Leong-Skomickova)

neglecta should be considered Endangered (EN D) in Singapore and at least Vulnerable
(VU Blab(iii)) globally (for details see Niissalo et ah, 2014).

Notes. The first collection of Hanguana neglecta on Bukit Timah dates back to 1885
and has been re-collected from the same location several times since. It was Tillich
& Sill (1999) who first realised that this species is not Hanguana malayana. Without
adequate material to hand, they did not pursue a description, but clearly labelled this
species as Hanguana ‘Singapur’ in their morphological study. A complete description
of Hanguana neglecta and a list of specimens examined are provided in Niissalo et al.
(2014).

Hanguana nitens Siti Nurfazilah, Mohd Fahmi, Sofiman Othman & P.C. Boyce,
Willdenowia 40: 207 (2010). - TYPE: Peninsular Malaysia, Johor Bahru, Mersing,
Hutan Simpanan Lenggor, 2°1572.7"N, 103°4376.7"E, 55 m, 18 April 2010, Siti
Nurfazilah bt Abdul Rahman, Boyce, P.C. & Ooi Im Hin HA-48 (holotype KEP; not
yet deposited as of 5 September 2014). (Fig. 4, 5)

Ecology and distribution. Lowland humid swamp forests, in blackwater mires or slow
moving streams. So far known to occur in Peninsular Malaysia and Singapore.

Hanguana in Singapore

9

Fig. 4. Hanguana nitens Siti Nurfazilah, Mohd Fahmi, Sofiman Othman & P.C.Boyce. A.
Habit. B. Detail of ripe fruits (inset upper left: male flower buds; inset lower left: young
unripe fruits); note the separate, erect, pointed stigma lobes. C. Infructescence. From native
population at MacRitchie sector. Central Catchment Nature Reserve, JLS-3029. (Photos: Jana
Leong- Skomicko va)

10

Gard. Bull. Singapore 67(1) 2015

Fig. 5. Hanguana nitens Siti Nurfazilah, Mohd Fahmi, Sofiman Othman & P.C.Boyce. A.
Detail of tepals (side view). B. Detail of inner tepal, staminode and staminodial seale. C. Detail
of stigma on young fruit. D. Cross seetion of fruit showing single seed and two empty locules.
E. Longitudinal section of two-seeded fruit. F. Detail of stigma on fully ripe fruit. G. Seed (top
view). H. Seed (side lateral view); note there are no appendages along the incurved rim. I. Seed
viewed from bottom. From native population at MacRitchie sector, Central Catchment Nature

v

Reserve, JLS-3029. (Photos: Jana Leong-Skornickova)

Provisional lUCN conservation assessment. Although when first described Hanguana
nitens was known to occur in three locations in Johor, a revision of herbarium material
at SING and KEP suggests that this species is more widespread, with specimen records
from Pahang, Selangor and Perak. Based on an EOO, which is larger than 20,000 km^’
this species can be provisionally placed into category of Least Concern (LC) following
the lUCN criteria (lUCN, 2012). Future explorations should, however, focus on an
assessment of population sizes and potential decline in extent and/or quality of habitat

Hanguana in Singapore

11

and adjust accordingly with more accurate data. In Singapore, Hanguana nitens is
so far known from a single locality in the Central Catchment Nature Reserve with
an area of occupancy (AGO) of c. 200 m^ and a predicted continuing decline in
quality of habitat. The colony of about 150 full-sized shoots consists of both male and
female plants, although considering the stoloniferous colonial nature of Hanguana
nitens, it almost certainly represents only a few individuals (genotypes), and as such
the species should be considered Critically Endangered (CR D) at the national level
(Davison et ah, 2008). Conservation should focus on regular harvesting of ripe fruits,
re-introduction into suitable habitats, and establishment and propagation of material to
saturate potential horticultural interest.

Additional specimens examined. SINGAPORE: Bukit Panjang: 1901, Ridley, H.N.1397
(SING); ibidem, 1906, Ridley, H.N. s.n. (SING); Bukit Timah Road: 19 Dec 1900, Ridley,
H.N. sji. (SING); Central Catchment Nature Reserve: MacRitchie sector, 21 May 2014,
Leong-Skornickovd, J. & Thame, A. JLS-3028 (SING); ibidem, 1 Jul 2014, Leong-Skornickovci,
J. & Thame, J. JLS-3029 (SING); Changi: 4 Oct 1890, Ridley, H.N. s.n. (SING); Chua Chu
Kang [Choa Chu Kang]: 1905, Ridley, H.N. s.n. (SING); With no precise locality: Feb 1837,
Gaudichaud, M. 112 (P, 3x).

PENINSULAR MALAYSIA: Johor: sine dat, Best 8290 (SING); Batu Pahat, Ayer Hitam
F. R., 21 Nov 1965, Ng KEP 100008 (KEP); Ponitan, Pengkalan Raja, sine dat., Alvins 72
(SING); ibidem, 28 Jun 1939, Ngadiman 36651 (SING); Kota Tinggi, Nam Heng estate, 20
Mar 1926, Teniya 640 (SING). Pahang: Kuantan, Sg. Baging F. R., 6 Oct 1989, Khairuddin
FRI 35418 (KEP, 2 sheets). Selangor: Petaling, Batu Tiga, Feb 1904, Ridley, H.N. s.n. (SING,
2 sheets). Kuala Lumpor, 1890, [? Ridley] s.n. (SING); Klang, Changgang, 3 Oct 1937, Mohd
Nurs.n. (SING). Perak: Hilir Perak, Hutan Melintang F.R., 13 Sep 1967, Ng FRI 5698 (KEP);
Taiping, BatuTogoh, Jun 1888, Wray 2128 (SING).

Note: Specimens of Hanguana nitens from Johor cited by Siti Nurfazilah et al. (2010) have not
yet been deposited at KEP as of 5 September 2014.

Notes. The original description of Hanguana nitens as given by Siti Nurfazilah et al.
(2010) largely agrees with our observation of the Singapore population, although the
plants in Singapore are overall more robust, reaching 2 m in height. The largest leaves
reach up to 2.2 m in length with leaf blades to 115 x 1 3.5 cm. Female inflorescences/
infiiictescences are also more robust with the peduncle and rachis reaching 1.1 m
and the median branches on the lowermost levels reaching up to 27 cm in length.
Male inflorescences were not included in the original description of the species (Siti
Nurfazilah et al., 2010). We observ^ed young male inflorescences in May, but with as
yet unopened buds. The structure of the inflorescences is similar to the female plants,
but generally much more slender. Flower buds appear in dense clusters of four to
seven. The tepals are bright green with a bronze tinge externally (Fig. 4B - inset upper
left).

Hanguana nitens certainly has ornamental potential with its beautiful glossy
corrugate leaves which, especially in young leaves, exhibit a pattern of mid-green and
dark green (Fig. 4). Its landscape usage may be similar to Hanguana malayana (as
cuiTently applied) but, as observed in the field, H. nitens is better suited to partially
shaded areas where the largest individuals occur, compared to H. malayana, which
thrives best in full sun.

12

Gard. Bull. Singapore 67(1) 2015

Hanguana mbinea Skomick. & P.C. Boyce, sp. nov.

Close to Hangtiana pantinensis in fruits ripening to ruby-red, but differing by a
more compact infructescence composed of shorter, stiffer partial inflorescences
attached almost peipendicularly to rachis, median branches to 11 cm with lateral
branches progi'essively shorter, and basally connate stigma lobes (compared to partial
infructescences composed of slender markedly erect branches to 14 cm long, median
and lateral branches of almost same length, and free stigma lobes). Hanguana mbinea
is also similar to H. triangulata (described below), but differs by fruits ripening to
ruby-red and stigma lobes connate basally (sometimes imperfectly) with free round
apices forming bluntly triangular structure (vs ripe fruits cream-white and stigma lobes
with sharply acute apices, connate at base, forming a sharply equilaterally triangular
structure in H. tiiangulata). - TYPE: Singapore, Central Catchment Nature Reserve,
forest around Upper Seletar Reservoir, 19 August 2014, Leong-Skornickovd, J. &
Thame, A. JLS-3037 (holotype SING; isotypes E, KEP). (Fig. 6, 7)

Herbaceous^ dioecious mesophyte to 1.5 m tall; stem terete, to 2.5 cm in diam.,
basally semi-ascending, with age becoming leafless at base, terminally ascending
with crown of up to 25 leaves; stolons absent. Leaves to 165 cm long, spreading
then arching, bases imbricate with hyaline margins (young leaves), turning erose-
marcescent with age; pseudopetiole 50-85 cm long, c. 13 mm wide, accounting
for 1/3-1 /2 of entire leaf length, roundly channelled with sharp margins; leaf blade
65-95 X 12-16 cm, narrowly elliptic, base attenuate, tip long and narrowly attenuate
with apicule 1 5-20 mm, leathery, adaxially mid to dark green, sparsely hairy (silky
appressed hairs evenly distributed; less visible in older leaves), abaxially lighter
green when ifesh, sparsely with silky appresed hairs evenly distributed (denser than
on upper surface); midrib weakly impressed, of the same colour as the rest of the
lamina adaxially, round-raised, mid green, almost glabrous and shiny abaxially.
Male inflorescences not observed, female inflorescences erect at anthesis, of same
structure and dimensions as infructescence. Infructescence erect, comprising up to
8 partial, whorled, altemate-secund, thyrsoid infructescences plus a tenninal spike;
partial infructescences spreading almost perpendicularly to rachis; peduncle and
rachis together up to 50 cm tall, dark purple-brown when fresh, conspicuously pale
brown-grey flocculose, visible portion of peduncle up to 20 cm long; one sterile bract
per peduncle, foliaceous, persistent, narrowly ovate with a basal claw, 88 (inch 8
cm long claw) x 14.5-11.5 cm; bract subtending partial infructescences similar to
sterile bracts, the bract supporting most basal partial infructescence c. 38 x 9 cm,
diminishing in size distally along the infructescence and fully reduced in uppermost
partial infructescences; partial infructescences each comprising up to 11 branches at
basal levels (occasionally two branches connate at base), fewer towards the apex of
the inflorescence, branches arising simultaneously from the axil of the subtending
bract, lateral branches progressively shorter in length (outermost lateral branches c.
y4 of the median branch), median branches at basal levels usually further branched
7-11 cm long, 3-4 mm in diam. Female flowers scattered, always solitary, sessile, all
with an associated minute bract and bracteole; perianth composed of 6 tepals in two

Hanguana in Singapore

13

Fig. 6. Hanguana rubinea Skomick. & P.C.Boyce. A. Habit. B. Detail of rachis with prominent
dense floeeulose indumentum, whieh is easily rubbed off (inset: detail of fruits and stigma). C.
Infructescenee. From type (Photos: Jana Leong-Skornickova)

14

Gard. Bull. Singapore 67(1) 2015

Fig. 7. Hanguana rubinea Skomick. & P.C.Boyce. A. Detail of tepals tightly clasping the base
of the fruit. B. Detail of inner tepals, staminodes and staminodial scales. C. Detail of stigma. D.
Cross section of fruit showing single seed and two empty locules. E. Longitudinal section of
fruit. F. Fruit in top view, after removal of apical half of the pulp, showing two seeds and one
empty locule; note the acute appendage at the top of the seed. G. Seed (top view). H. Seed (side
lateral view). I. Seed in semi-lateral view showing an acute appendage. From type JLS-3037.
(Photos: Jana Leong-Skomickova)

whorls tightly clasping ovary/fruit in fresh material, all tepals with prominent bulbous
thickening at base (more prominent in outer whorl), light green with more or less dense
minute red-brown speckles, margin c. 0.2 mm wide, hyaline translucent white; outer
tepals semi-eircular, 1.5-2. 5 mm long, c. 2.5 mm broad, connate at base, sparsely
araehnoid; inner tepals almost semi-circular, 3-3.5 mm long, 4^.5 mm broad at base,
free to base, almost glabrous (occasionally sparsely arachnoid); staminodes 6, in two
whorls, pale green to eream white, triangular, outer staminodes, 0.5 mm long, 0.5 mm

Hanguana in Singapore

15

broad at base, inner staminodes longer, c. 1 mm long, 0.5 mm at base, each basally
sheathed with a broad narrow scale (often shallowly bilobed), c. 0.5 mm long, and c.
2 mm broad, brown with translucent margin; ovary green, ovoid, glossy green; stigma
3-lobed, each lobe 1.2-1. 4 mm long (fruiting material), broadly ovate (-to bluntly
trullate) with round apex, lobes connate basally (sometimes imperfectly) with free
apices forming bluntly triangular structure (c. 2,5 mm in diam, in fruiting material)
with points of connation seen as grooves, green (flowering stage), matte dark brown
(fruiting stage). Ripe fruit globose, 9-10 imn diam., dark pink-red externally, pulp 1-2
mm thick, cream- white, fairly hard, exuding yellow juice when disturbed, ripenmg
from bright green through cream- white to dark pink-red; seeds 1 -2 per fruit, c. 5 x 4
mm, brown, bowl-shaped with slightly incurved margins, with a triangular appendage
positioned on the distal part of the rim, c. 5 x 4 mm, deeply excavated, cavity filled
with placental tissue.

Etymology. The specific epithet is derived from the ruby-red colour of the ripe fruits.

Ecology and distribution. Growing on the slopes of, or in the proximity of, small/
seasonal streams in primary or partially disturbed primary lowland forest. So far
endemic to Singapore.

Provisional lUCN conservation assessment. Based on our recent collections and on
reliably identified fertile herbarium records from the past 30 years, Hanguana ruhinea
occurs in four locations in Singapore (Bukit Timah, Mandai, MacRitchie and Seletar).
The extent of occun*ence (EOO) is c. 12 km-, the habitat is fragmented and, based on
our observations, the number of adult individuals is fewer than 250, with fewer than
50 individuals in each sub-population. With impending and proposed developments
in some of the existing locations, and serious damage caused by wild boars observed
in two locations, further decline of the populations and further fragmentation of the
habitat is foreseen. Hanguana rubinea should be therefore considered as Critically
Endangered (CR C.2) locally (Davison et al., 2008), and because it is so far endemic
to Singapore, also globally CR Blab(iii,v); C2a(i)) (lUCN, 2012).

Additional specimens examined. SINGAPORE: Bukit Timah Nature Reserve: upper end of
Taban Loop, 28 May 2014, Leong-Skornickovd, J. & Thame, A. JLS-2791 (SING); Central
Catchment Nature Reserve: sector 53 [MacRitchie] 29 Apr 1992, Yong et al. NSR 598
(SING); MacRitchie, off Shinto Trail, 1 Jiil 2014, Leong-Skornickovd, J. & Thame, A. JLS-
3027 (SING); MacRitchie, off Lomie Trail, 14 Aug 2014, Leong-Skornickovd, J, & Thame, A.
JLS-3030 (SING); Chan Chu Kang: 14 Apr 1890, Goodenough, J.S. s.n. (SING); Mandai:
Mandai Forest, 6 Jan 2009, Gwee, A.T SING 2009-09 (SING); Mandai forest, 19 Aug 2014,
Leong-Skornickovd, J. & Thame, A. JLS-3034 (SING; sterile); Ponggol: 1905, Ridley, H.N.
s.n. (SING).

Note: The following two specimens might also represent ifr/ngwana rubinea, but the specimens
deviate slightly from typical H. rubinea and are cited here with caution: Mandai Road, 28 Jul
1929, Corner, E.J.Hs.n. (SING); Seletar, 29 Mar 1889, Ridley, H.N 170 (SING).

16

Gard. Bull. Singapore 67(1) 2015

Notes. In late fmiting stage, Hanguana rubinea is easy to recognise by its pretty
ruby-red fruits. In early fruiting stages, when the ovaries are still creamy white, it
can potentially be mistaken for Hanguana tnangulata with which it shares a similar
infructescence stmcture with partial infmctescence branches attached almost
perpendicularly to the rachis. However, the shape of the stigma is unmistakeable in the
latter species as the three stigmatic lobes with acutely sharp apices fonn an equilateral
triangle (compare Fig. 7C and 9C). Further differences are in the leaves and the seeds.
The leaf blades of Hanguana rubinea are almost flat, or only weakly corrugate, with
an evenly distributed abaxial silky indumentum, compared to H triangulata which has
corrugated leaves and the indumentum is more visible and flocculose. While the seeds
of both species are bowl-shaped, Hanguana rubinea has an acute appendage along
the rim (Fig. 7G-1) compared to a blunt appendage composed of two sub-lobes in H
triangulata (Fig. 9G-1).

V

Hanguana triangulata Skomick. & P.C.Boyce, sp. nov.

Similar to Hanguana exultans by fmits ripening cream-white to pale yellow, but
differing by a more compact infructescence composed of shorter, stiffer partial
inflorescences attached almost perpendicularly to rachis and stigma lobes with sharply
acute apices, connate at base, fonning an equilateral triangular structure (vs partial
infructescence somewhat erect, attached at c. 45° to rhachis and stigma composed
of three free lobes). - TYPE: Singapore, Bukit Timah Nature Reserve, slopes around
Taban Loop, 28 May 2014, Leong-Skornickovd, J. & Thame, A. JLS-2789 (holotype
SING). (Fig. 8, 9)

Herbaceous, dioecious mesophyte to c. 1 .6 m tall; stem terete, to 2.5 cm in diam., basally
semi-ascending, with age becoming leafless, terminally ascending with crown of up 20
leaves; stolons absent. Leaves to 150 cm long, spreading then arching; bases imbricate,
margins hyaline (young leaves), turning erose-marcescent with age; pseudopetiole
40-50 cm long, c. 12 mm wide, accounting for c. 1/3 of entire leaf length, roundly
channelled with sharp margins, sparsely softly flocculose; leaf blade 90-106 x 12-13
cm, nan*owly elliptic, base attenuate, tip long and narrowly attenuate with apicule
to 5 mm, leathery, adaxially dark green, sparsely hairy (silky appressed hairs evenly
distributed; less or not visible in older leaves), abaxially mid-green, somewhat shiny,
covered with silky flocculose hairs (unevenly distributed, much denser than on upper
surface); midrib weakly impressed, almost of the same colour as the rest of the lamina,
sparsely flocculose (especially towards the base) adaxially, round-raised, mid green,
sparsely flocculose and shiny abaxially. Female and male inflorescences not observed,
although, based on observations of infructescence architecture, almost certainly erect
at anthesis. Infructescence erect, comprising up to 6 partial, whorled, alteraate-secund,
thyrsoid iiifructescences plus a terminal spike; partial infructescences spreading
almost perpendicularly to rachis; peduncle and rachis together up to 70 cm tall, green
when fresh, conspicuously pale brown-grey flocculose, visible portion of peduncle
up to 40 cm long; sterile bracts two per peduncle, foliaceous, narrowly ovate with a

Hanguana in Singapore

17

Fig. 8. Hanguana triangulata Skornick. & P.C.Boyce. A. Habit. B. Detail of fruits and the
typieal sharply triangulate stigma. C. Inflorescence. From type 759. (Photos: Jana Leong-
Skornickova)

18

Gard. Bull. Singapore 67(1) 2015

Fig. 9. Hanguana triangulata Skomick. & P.C.Boyce. A. Detail of tepals (side view). B. detail
of inner tepals, staminodes and staminodial scales. C. Detail of stigma. D. Cross section of
fruit showing single seed and two empty locules. E. Longitudinal section of fruit. F. Fruit in
basal view, after removal of tepals, showing three staminodial scales still attached. G. Seed
(top view). H. Seed (side lateral view). I. Seed (semi-lateral view showing a blunt appendage
composed of two sub-lobes. From type JLS-2789. (Photos: Jana Leong-Skomickova)

basal claw, persistent, 36-71 x 6.5-11.5 cm; bract subtending partial infructescences
similar to sterile bracts, the bract supporting most basal partial infmctescence 20
X 3 cm, diminishing in size distally along the infmctescence and fully reduced in
uppermost partial infructescences; partial infructescences each comprising up to 10
branches at basal levels (fewer towards the apex of the inflorescence), branches arising
simultaneously from the axil of the subtending bract, lateral branches progressively
shorter in length (outermost lateral branches 2/3- 1/2 of the median branch), median
branch 6-8 cm long, c. 3 mm in diam. Female flowers scattered, always solitary.

Hanguana in Singapore

19

sessile, all with an associated minute bracteole; perianth composed of 6 tepals
in two whorls tightly clasping ovary/fruit in fresh material, outer 3 tepals broadly
triangular with round apex, all tepals with prominent bulbous thickening at base (more
prominent in outer whorl), light green, margin c. 0.2 mm wide, hyaline translucent
white; outer tepals semi-circular, c. 2 mm long, c. 2.8 mm broad, connate at base,
sparsely arachnoid; inner tepals almost semi-circular, c. 3 mm long, c. 4 mm broad,
free to base, almost glabrous (occasionally sparsely arachnoid); staminodes 6, in two
whorls, cream-white, triangular to naiTowly triangular, outer staminodes minute, c.
0.3 mm long, 0.2 mm broad at base, inner staminodes larger, c. 1 mm long, 0.3 mm at
base, each basal ly sheathed with semi-circular staminodial scale, c. 0.8 mm long, and
c. 1.3 mm broad, brown with irregular (usually bilobed) translucent margin; stigma
3-lobed, lobes connate basally, each lobe c. 1.5 mm long (fruiting material), trullate
with shaiply acute apex, lobes perfectly connate basally forming equilateral triangle
(c. 2.5 mm in diam. in fruiting material), raised, matte dark brown in late fruiting stage.
Ripe fruit cream externally, pulp c, 2 mm thick, hard, cream-white, exuding yellow
juice when disturbed, globose, 9-10 mm diam., ripening from bright green to cream-
white; seeds usually two per fruit, c. 5-6 x 4.5-5 mm, brown, broadly boat-shaped to
bowl-shaped, deeply excavated, with a blunt appendage composed of two sub-lobes
positioned on the distal part of the rim, cavity filled with placental tissue.

Etymology. The specific epithet refers to the sharply triangular shape of the stigma, a
character which is conspicuous even in herbarium material.

Ecology and distribution. Hanguana triangulata is an undergrowth species growing
in the proximity of streams in lowland evergreen forest. So far endemic to Singapore.

Provisional lUCN conservation assessment. Only two locations in Singapore with
fewer than 20 individuals in all (AOO 0.5 km2) were observed during recent fieldwork.
There are signs of severe damage caused by wild boars in some adult individuals,
thereby warranting a status of Critically Endangered (CR D) at the national and, given
its endemic status, also global levels (Davison et al., 2008; lUCN, 2012). Immediate
conservation efforts to ensure the survival of this species have focused on the harvesting
of ripe fruits, optimising the protoeol for ex situ cultivation and multiplication, and re-
introductions to suitable habitats.

Additional specimens examined. SINGAPORE: BukitTimah: Ridley, H.N. 9531 (SING);

Bukit Timah Nature Reseive, slopes around Taban Loop, 12 Aug 2014, Leong-Skornickovd, J.
& Thame, A. JLS-2792 (SING); ibidem, slopes around Tiup Tiup path, 14 Sep 2014 , Leong-
Skornickovd, J. J LS- 3 04 6 (SXHGy, Central Catchment Nature Reserve: forests around Upper
Seletar Reservoh, 19 Aug 2014, Leong-Skornickovd, J. & Thame, A. JLS-3036 (SING, sterile);
Kranji: 5 Aug 1889, Goodenoiigh, J.S. s.n. (SING); Pulaii Ubin: Aug 1898, Ridley, H.N. s.n.
(SING).

Notes. Hanguana triangulata is currently known to occur only in the Bukit Timah
Nature Reserve (confirmed from fruiting material) and in the Central Catchment

20

Gard. Bull. Singapore 67(1) 2015

Nature Reserve in forests around Upper Seletar Reservoir (identification based on
sterile material only). Historical herbarium records suggesting its past presence in
Kranji and Pulau Ubin are both over a century old. The sharply triangular stigma
is easily observable in dried material, although the sharp tips may cuiTe somewhat
downwards. Seeds extracted from material as old as 1 889 match well to seeds extracted
from recently collected material in Bukit Timah Nature Reserve (both exhibiting the
blunt weakly bilobed appendage, see Fig. 9G-I).

Introduced and naturalised species

V

Hanguana corneri Skomick. & RC.Boyce, sp. nov.

Unique amongst cuiTently known Hanguana species by its leaves which are shiny
emerald-green above and deep purple and silvery flocculose beneath. - TYPE:
Peninsular Malaysia, Johor, Sungai Berassau, Mawai-Jemulang Road, 7 February
1935, Corner, E.J.H.s.n. (holotype SING!). (Fig. 10)

Herbaceous, dioecious mesophyte to c. 0.8 m tall; stem terete, to 2 cm in diam.,
basally semi-ascending, with age becoming leafless at base, terminally with crown
of 15-20 leaves; stolons absent. Leaves to 85 cm long, spreading then arching, bases
imbricate with hyaline margins (young leaves), turning erose-marcescent with age;
pseudopetiole 10-20 cm long, accounting for !4-‘/3 of entire leaf length, roundly
channelled with sharp margins; leaf blade 54-66 x 10-14.5 cm, narrowly elliptic
with undulate margins, base attenuate, tip nan'owly acute with apicule 2-3 mm, softly
leathery, adaxially dark green, with glossy lustre, abaxially purple red, with dense soft
appressed flocculose indumentum; midrib weakly impressed, mid-green (much lighter
than the rest of the lamina) adaxially, round-raised, light pink-brown, flocculose
abaxially. Female inflorescence not observed, although, based on obseiwations of
infmctescence architecture, almost certainly erect at anthesis. Male inflorescence
erect at anthesis, comprising 6 partial inflorescences and a terminal spike; peduncle
and rachis c. 30 cm long; peduncle c. 6 cm long, no sterile foliaceous bract observed;
bract subtending lowest partial inflorescence foliar, naiTOwly ovate, c. 19 cm long
(width obscured in type specimen), with basal claw c. 2 cm, distally diminishing in size
to narrowly triangular bracts, fully reduced at uppermost partial inflorescences. Male
flowers scattered, in groups of (2-)3-5; perianth composed of 6 tepals in two whorls,
all tepals ovate, greenish (according to a note on type specimen), outer tepals c. 1 mm
long, inner tepals c. 1 .5 mm long (measured from dried flower buds). Female flowers
scattered, in pairs or solitary, sessile, all with an associated minute bract and bracteole;
perianth composed of 6 tepals in two whorls tightly clasping fruit (in dry material),
all tepals with prominent bulbous thickening at base (more prominent in outer whorl);
outer tepals semi-circular, c. 1 mm long, c. 1.5 mm broad, connate at base; inner
tepals almost semi-circular, c. 2 mm long, 2-2.5 mm broad, free to base; staminodes
and staminodial scales not observed; stigma 3-lobed, each lobe c. 7 mm long (dried
fruiting material), ovate with blunt apex, lobes perfectly connate basally with free

Hanguana in Singapore

21

Fig. 10. Hanguana corneri Skornick. & P.C.Boyce. A. Habit. B. Base of the plant showing
semi-aseending leafless stem. C. Detail of lamina abaxially, showing the unique dark purple-
red eoloration and dense cover of silvery flocculose hair. From plant growing at Bukit Timah

V

Nature Reserve, JLS-2790. (Photos: Jana Leong-Skomickova)

22

Gard. Bull. Singapore 67(1) 2015

apices forming bluntly triangular structure with raised centre (c. 1.1 mm in diam., in
dried fruiting material), matte dark brown in late fruiting stage. Infmctescence erect,
comprising up to 5 partial altemate-secund, thyrsoid infructescences plus a teraiinal
spike; partial infructescences semi-erect, at an angle c. 40°-50“ to rachis; peduncle
and rachis together up to 45 cm tall; peduncle up to 10 cm long, one sterile bract per
peduncle, foliaceous, persistent, nari'owly ovate with basal claw, c. 35 (inch 5 cm claw)
X c. 6 cm; bract subtending partial infructescences similar to sterile bract, distally
diminishing in size to nan'owly ti'iangular bracts, fully reduced in uppermost partial
inflorescences; partial infructescences each comprising up to 5 branches at basal levels,
fewer towards the apex of the inflorescence, branches arising simultaneously from the
axil of the subtending bract, lateral branches progressively shorter in length (outemiost
lateral branches c. % of the median branch), both median and lateral branches usually
further branched, median branch to 10 cm long, c. 2.5 mm in diam. Fruit (described
from old dried fruit) globose, c. 4 mm in diam.; seeds brown, c. 3-3.5 mm in diam.,
34 globose to ellipsoid, ostiole wedge-shaped accounting for c. % of the seeds, deeply
excavated, cavity filled with placental tissue.

Etymology. We name this species for E.J.H. Comer (1906-1996), a botanist and
mycologist who first collected this beautiful species. He was a giant among tropical
Asian botanists and, in particular, for ‘Malayan’ botany.

Ecology and distribution. Native to Peninsular Malaysia (Johor) and introduced
to Singapore. According to the notes on Comer’s specimen, this species occurs in
swampy forests with flowering occuming in Febmary. Populations of what is almost
certainly the same species, although not occurring in swampy areas, are known from
SW Sarawak (Boyce, pers. obs.), notably in the Penrissen Range, where plants occur
intermixed with another two as-yet undescribed Hanguana species a few metres from
a precipitous escarpment marking the border with Kalimantan Barat.

Provisional lUCN conservation assessment: Owing to a lack of data on the distribution
and population sizes of this species in Peninsular Malaysia, and the uncertainty as to
whether this species also extends to Borneo, we propose that this species be treated as
Data Deficient (DD).

Additional specimens examined. SINGAPORE: Bukit Timah Nature Reserve: slopes around
Taban Loop, 28 May 2014, Leong-Skornickovd, J. & Thame, A. JLS-2790 (SING).

Notes. A single plant of this conspicuous species has been discovered in the Taban
Loop area of Bukit Timah Nature Reserve (BTNR). A revision of material from
Peninsular Malaysia at SING revealed the existence of a male flowering specimen
of this species, collected by E.J.H, Comer in 1935 from Sungai Berassau (Peninsular
Malaysia, Johor). Although it is almost certain that living plants of this species were
brought to Singapore from a field trip conducted by Comer, who at that time worked
in Singapore Botanic Gardens (1929-1945), it is not clear how they ended up on Bukit

Hanguana in Singapore

23

Timah. The first author has observed, as part of earlier work on native Zingiberales,
that several Scaphochlamys species, all of them certainly not native to Singapore, also
occurred around the same area. Some of these Scaphochlamys species were described
as new to science by R.E. Holttum from herbarium material collected by Comer.
Holttum (1950) explicitly stated that he based his descriptions solely on the herbarium
material and that he had never seen these species alive, making any connection of
Holttum to the Bukit Timah plantings unlikely. According to John Dransfield and Ruth
Kiew, who both pursued their PhD studies under Comer’s supervision and knew him
well, it is also highly unlikely that it was Comer himself who planted these on Bukit
Timah (J. Dransfield, R. Kiew, pers. comm.). While there are no official records, we
tend to agree with a suggestion by Dransfield, that living plant material was most
likely brought back to Singapore Botanic Gardens by Comer’s field assistants Kiah
and Md. Nur. When and by whom the material was planted on Bukit Timah, however,
remains unclear.

Several plants of Hanguana corneri are cultivated in the living collections at
the Forest Research Institute, Malaysia. These collections, unfortunately, lack exact
provenance record, although they are also likely to be from Johor (Saw Leng Guan,
pers. comm.).

The above description is based on cuirently available material consisting of
sterile living plants m the Bukit Timah Nature Reserve, a male dried specimen (the
type), and old infmctescences observed in the living collections of the Forest Research
Institute, Malaysia. While the description is incomplete, we feel that the species is
so distinct and recognisable, even in herbarium specimens, that it wamants fomial
description. The description can be further improved with additional material gathered
in the future.

Hanguana malayana (Jack) Merr., Philipp. J. Sci., C, 10: 3 (1915). - Veratrum
malayanum Jack, Malayan Misc. 1(1): 25 (1820). - Veratronia malayana (Jack) Miq.,
FI. Ned. Ind. 3: 553. (1859). - Susum malayanum (Jack) Planch, ex Hook, f, FI. Brit.
India 6: 391 (1892). - TYPE: Malaysia, Pulau Pinang [“Poeloe Pinang”], (not

traced). (Fig. 11, 12)

Hanguana anthelminthica (Blume ex Roem. & Schult.) Masam., Enum. Phan. Bom.:
81. (1942). — Susum anthelminthicum Blume ex Roem. & Schult., Syst. Veg. 7(2):
1493 (1830). — Susum malayanum f. aquatica Backer, Handb. FI. Java 3: 3 (1924).
— Hanguana malayana subsp. anthelminthica (Blume ex Roem. & Schult.) Backer,
Bekn. FI. Java 10(212): 2 (1949). — Hanguana malayana var. anthelminthica (Blume
ex Roem. & Schult.) Bakli., Blumea 6: 399 (1950). - TYPE: Indonesia, Java “in
paludibus circa Buitenzorg”, Blume s.n. (lectotype L! [L21 07638, L21 07639; both
sheets cross-referenced as being part of single gathering], here designated; isotype BO
n.v. (as per Siti Nurfazilah et al., 2010).

Hanguana aquatica Kaneh., Trans. Nat. Hist. Soc. Formosa 25: 8 (1935). - TYPE:
Caroline Islands, Palau, Almonogni, Babeldaob, 13 April 1938, S. Hatusima 4866
(holotype FU!).

24

Gard. Bull. Singapore 67(1) 2015

Fig. 11. Hanguana malayana (Jack.) Merr. A. Habit. B. Branches with ripe fruits (inset: close-
up detail of fruits and stigma). C. Infrueteseenee. From plants eultivated at Singapore Botanie

v

Gardens, JLS-3033. (Photos: Jana Leong-Skornickova)

Hanguana in Singapore

25

Fig. 12. Hanguana malayana (Jack.) Merr. A. Detail of tepals clasping base of ripe fruit (side
view). B. Detail of inner tepals, staminodes and staminodial scales. C. Detail of stigma. D.
Cross section of fruit showing three well-developed seeds. E. Longitudinal section of fruit. F.
Detail of tepals clasping base of ovary (side view). G. Seed (top view). H. Seed (bottom view).
I. Seed (lateral view). From plants cultivated at Singapore Botanic Gardens JLS-3033. (Photos:
Jana Leong-Skornickova)

Notes: The identity of Hanguana malayana has been the subjeet of mueh debate, as
the original material by Jack is still missing and Jack’s original description, based
on material from Pinang [Pulau Penang], although fairly long, does not offer many
characters which could be considered species specific. The name has been uncritically
applied to almost all forest species in Malaysia and Singapore since Backer’s treatment
of Hanguana for Flora Malesiana (Backer, 1951). The most recent attempt to clarify
the identity and circumscription of Hanguana malayana is that of Siti Nurfazilah et al.
(2010), who proposed that the name H. malayana should be applied to a widespread

26

Gard. Bull. Singapore 67(1) 2015

large, helophytic, stoloniferous species. As this is the species which occurs in cultivation
in Singapore (Fig. 11, 12), this current application is followed here with caution.

Hanguana malayana (sensu Siti Nurfazilah et aL, 2010) is freely available in
Singapore’s nurseries and is occasionally used in landscaping. All herbarium records
originating from Singapore, and previously identified as Hanguana malayana, at the K
and SING herbaria (no specimens from Singapore exist at KEP), turned out to belong
to the four species above treated. Our field explorations have so far not confirmed the
presence of any populations which could be considered to be wild, although populations
of Hanguana malayana have been planted in various locations in Singapore where
they thrive and may naturalise easily. There is also no confirmed herbarium record
of Hanguana malayana from areas close to Singapore. A revision of all Hanguana
herbarium sheets at K, KEP, P and SING revealed that there are no collections of H.
malayana from Johor. All confinned collections in Peninsular Malaysia were collected
only in Perak, Selangor, Terengganu, and a single historical collection of Griffith 6014
(K), collected in Malacca [Melaka], which is about 200 km from Singapore. We have
located two collections of Hanguana malayana from Sumatra (Indonesia), of which
Yates 2281 (P, 4x) is from Medan (c. 600 km from Singapore), while Zollinger’s undated
collection is without precise locality. Considering the above, there is, therefore, no
reliable evidence, historical or recent, which would support the theory that Hanguana
malayana was ever native to Singapore.

Conclusion

Recent discoveries of six native species new to science in heavily urbanised Singapore
(including Hanguana rubinea and //. triangulata described above; H, neglecta -
Niissalo et ah, 2014; Zingiber singapurense Skomick. - Leong-Skomickova et ah,
2014; two Utania species - Sugumaran & Wong, 2014), serves to remind us of several
important points. Firstly, that although Singapore is the most densely collected country
in SE Asia and its primary vegetation has been severely disturbed, the process of
documenting and fully understanding its rich biodiversity is incomplete. Secondly,
that while herbarium-based taxonomy is sufficient in some plant groups, satisfactory
progress on others without observation of fertile material in the field is almost
impossible (for example in Zingiberaceae and Hanguanaceae). And lastly, that such
field-based knowledge can lead to the discovery of previously overlooked characters,
in this case the staminodial scales and the seed stmcture, which can then be applied to
the identification of existing herbarium collections.

A poor understanding of the genus Hanguana in Singapore and the misapplication
of the name Hanguana malayana, which was previously treated as Vulnerable in
the Singapore Red Data Book (Davison et al., 2008), obviously has implications
for conservation of all four native species. These four species are all currently
considered to be Endangered or Critically Endangered locally, and those endemic to
Singapore also globally {H. neglecta ENATJ, H. nitens CR/LC, H. rubinea CR/CR
and H. triangulata CR/CR). Further conservation work, including the observation of

Hanguana in Singapore

27

pollinators and dispersal agents and the DNA barcoding of native species is planned.
Also, now that the hidden diversity of Hanguana in Singapore has been recognised, in
situ and ex situ conservation and propagation efforts for this interesting yet extremely
neglected plant family in Singapore is underway. Due to the lack of any records
suggesting that the large stoloniferous helophytic species, currently called Hanguana
malayana, was ever native to Singapore, we recommend that this species be treated as
not native to Singapore.

ACKNOWLEDGEMENTS. The first author thanks the National Parks Board, Singapore.
Our sincere thanks are due to numerous NParks staff and volunteers, who provided us with
continuous support in our taxonomic and conservation research on the Singapore flora, in
particular: Desmond Lee, Mishak Sunari (Sungei Buloh Wetland Reserve, field support),
Derek Liew, Chew Pin Ting (Central Catchment Nature Reserve, field support), Ang Wee Fong
(Horticulture and Community Gardening Division), Aung Thame, Sunia Teo and Michael Leong
(Singapore Botanic Gardens, field support and propagation of native species for conservation
purposes). We thank Prof Hans-Juergen Tillich for fruitflil discussions and suggestions on the
possible origin and function of staminodial scales, and confinnation of the identity of Hanguana
bogneri, and Dr Saw Leng Guan from the Forest Research Institute, Malaysia for granting
access to FRIM’s living collections of Hanguana , which helped us improve the description
of the imperfectly known Hanguana corneri. We also thank Eva-Nicole Fisher (volunteer at
Singapore Botanic Gardens) for her help with a translation of the research paper by Tillich &
Sill (1999) which was originally published in German, and to the anonymous reviewers for
useful comments to improve this manuscript.

References

Airy Shaw, H.K. (1980). Anew species of Hanguana from Borneo. Kew Bull. 5(4): 819-821.

Bachman, S., Moat, J., Hill, A.W., de la Torre, J. & Scott, B. (2011 ). Supporting Red List threat
assessments with GeoCAT: Geospatial Conservation Assessment Tool. In: Smith, V. &
Penev, L. (eds) e-lnfrastmctures for data publishing in biodiversity science. ZooKeys
150: 117-126. (Version BETA)*

Backer, C.A. (1951). Hanguana. In: Steenis, C.G.G.J. van (ed) Flora Malesiana, ser. 1, 4(3):
248-250.

Bayer, C., Appel, O. & Rudall, P.J. (1998). Hanguanaceae. In: Kubitzki, K. (ed) The Families
and Genera of Vascular Plants, vol. IV. Pp. 223-225. Berlin-Heidelberg: Springer-
Verlag.

Beentje, H. (2012). The Kew Plant Glossary, an illustrated dictionary of plant terms (revised
edition). Royal Botanic Gardens, Kew: Kew Publishing.

Chong, K.Y., Tan, H.T.W. & Corlett, R.T. (2009). A Checklist of the Total Vascular Plant Flora
of Singapore: Native, naturalised and cultivated species. Singapore: Raffles Museum of
Biodiversity Research, National University of Singapore.

Davison, G.W.H., Ng, P.K.L. & Ho, H.C. (eds) (2008) The Singapore Red Data Book:
Threatened plants and animals of Singapore, 2nd ed. Singapore: The Nature Society
(Singapore).

28

Gard. Bull. Singapore 67(1) 2015

Holttum, R.E. (1950). The Zingiberaceae of the Malay Peninsula. Gard. Bull Singapore 13:1-
249.

lUCN (2012). lUCN Red List Categories and Criteria: Version 3.1. Second edition. Switzerland,
Gland and UK, Cambridge: TUCN.

Keng, H., Chin, S.C. & Tan, H.T.W. (1998). The Concise Flora of Singapore, volume IT.
Monocotyledons. Singapore: Singapore University Press.

Leong-Skomickova, J., Thame, A. & Chew, P.T. (2014). Notes on Singapore native Zingiberales
I: A new species of Zingiber and notes on the identities of two further Zingiber taxa.
Card. Bull. Singapore 66(2): 153-167.

Low, Y.W., Leong, P.K.F., Tee, S.P., Rajesh Singh, Tay, M.L.C. & Wong, K.M. (2014).
Margaritaria (Phyllanthaceae), a new generic record for the Singapore flora. Gard.
Bull. Singapore 66(1): 47-56.

Mohd Faluui, A.B., Alimad Sofiman, O. & Boyce, PC. (2012). Studies on Hanguana
(Commelinales: Hanguanaceae) for Sunda TV: Hanguana major clarified, a new species
from Sarawak, Malaysian Borneo, and description of male reproductive organs in H.
bakoensis. Ke\v Bull. 67: 205-211.

V

Niissalo, M., Wijedasa, L., Boyce, RC. & Leong-Skomickova, J. (2014). Hanguana neglecta
(Hanguanaceae): a new plant species from a heavily collected and visited reserve in
Singapore. 188(1): 14-20.

Rodda, M., Tran, H.D., Chiew, P.T., Liew, D. & Leong-Skomickova, J. (2012). The rediscovery
of Dischidia hirsuta (Apocynaceae, Asclepiadoideae) in Singapore. Gard. Bull.
Singapore 64(2): 293-299.

Rudall, PJ., Stevenson, D.W. & Linder, H.P. (1999). Stmcture and systematics of Hanguana, a
monocotyledon of uncertain affinity. Austral Syst. Bot. 12: 311-330.

Siti Nurfazilah, A.R., Mohd Fahini, A.B., Ahmad Sofiman, O. & Boyce, PC. (201 0). Studies on
Hanguana (Commelinales-Hanguanaceae) for Sunda II: Five new forest species, a new
species record from Peninsular Malaysia, and a redefinition of the taxonomic limits of
Hanguana malayana. Willdenowia 40: 205-219.

Siti Nurfazilah, A.R., Ahmad Sofiman, O. & Boyce, PC. (2011). Studies on Hanguana
(Commelinales-Hanguanaceae) for Sunda I: Hanguana bakoensis, a new forest species
from Sarawak, Malaysian Borneo, and notes on critical morphologies for elucidating
Hanguana tAxomvny. Acta Phytotax. Geobot. 61(3): 139-143.

Sugumaran, M. & Wong, K.M. (2014). Studies in Malesian Gentianaceae, VI. A revision of
Utania in the Malay Peninsula with two new species. PI. Ecol Evol. 147(2): 213-223.

Tillich, H.-J. (1996). Seeds and seedlings in Hanguanaceae and Flagellariaceae
(Monocotyledons). Sendinera 3: 187-197.

Tillich, H.-J. & Sill, E. (1999). Systematische studien zur morphologic und anatomic von
Hanguana Blume (Hanguanacae) und Flagellaria L. (Flagellariaceae). Mit der
Beschriebung einer neuen Art, Hanguana bogneri spec. nov. Sendtnera 6: 215-238.

Gardens’ Bulletin Singapore 67(1): 29-33. 2015
doi: 10.3850/S238258 12 15000022

29

Dischidia acutifolia (Apocynaceae, Asclepiadoideae)
a new record for the Singapore flora

M. Rodda\ D. Lee^ & S. Mishak^

^Herbarium, Singapore Botanic Gardens, National Parks Board,

1 Cluny Road, 259569 Singapore
rodda.michele@gmai] .com

^Conservation Division, Central Nature Reserve, National Parks Board,

1 Cluny Road, 259569 Singapore

ABSTRACT. We report the discovery of Dischidia acutifolia Maingay ex Hook.f., a new
record for the Singapore flora. The species is only known from a single locality in Sungei
Buloh Wetland Reserve and is therefore considered Critically Endangered in Singapore. A
lectotype for the name is selected.

Keywords. lUCN Red Lists, Sungei Buloh Wetland Reserve

Introduction

In Singapore there are ten native species of Dischidia R.Br. (Chong et al., 2009;
Rodda et ah, 2012). Five of these are considered nationally extinct and one, Dischidia
singaporensis Ridh, is considered a globally extinct species. A general introduction
to the genus Dischidia can be found in various recent local treatments (Rintz, 1980;
Li et ah, 1995; Forster et ah, 1996; Jagtap & Singh, 1999; Livshultz et ah, 2005)
while information on the local Singapore species is provided in Rodda et ah (2012). In
September 2013 Desmond Lee and Mishak Shunari found an unidentified Dischidia
growing at the base of the tmnk of Talipariti tiliaceum (L.) Fryxell in Sungei Buloh
Wetland Reserve. A small sterile cutting was brought to the Singapore Herbarium
for identification. The cutting was incorporated into the Singapore Botanic Gardens’
Research Living Collections where it rooted and soon after started flowering before
producing any new growth. It was then identified as Dischidia acutifolia, a species
rather common from Thailand to the Moluccas (Rintz, 1980). The species is not in
the most recent checklist of Singapore’s vascular plant flora (Chong et ah, 2009) and
no specimens collected in Singapore have previously been deposited in SING. We
therefore here report it as a new record for the Singapore flora.

We can speculate that the Singapore plant may originate from a recent
colonisation from seeds wind-dispersed from the coastal mangroves in Johore
(Malaysia). However it is equally likely that the species, an epiphyte often growing
high up in the tree canopy and bearing inconspicuous flowers, may have always been
part of the Singapore flora but was never previously collected.

30

Gard. Bull. Singapore 67(1) 2015

Conservation considerations

The species is only known from a single collection and is, therefore, to be considered
Critically Endangered in Singapore, according to guidelines given in The Singapore
Red Data Book (Davison, 2008). The original plant observed in 2013 was negatively
affected by the drought in early 2014 (Ziegler et ah, 2014), and the recovery has been
slow (Lee and Mishak, personal observation). In contrast, the clone is very vigorous in
cultivation and is being mass propagated at the Singapore Botanic Gardens (SBG ace.
no. 20132480), Stock will soon be released for reintroduction trials.

The morphological description below is based on the specimen collected in
Singapore. Measurements from cultivated materials are indicated in parentheses when
they are significantly different from wild material.

Taxonomy

Dischidia aciitifolia Maingay ex Hook.f, FI. Brit. India 4(10): 51 (1883). - TYPE:
Peninsular Malaysia, Malacca, 30 Oct 1 865 or 6, Maingay, A.C. 1960 {Kew distribution
number 1122) (lectotype K, designated here [K0009 11032]). (Fig. 1)

Plants succulent, herbaceous, glabrous; epiphytic or hemi-epiphytic vines growing
loosely rooted on the host tree stems, clinging by adventitious roots, rarely pendulous.
Latex white. Roots on mature plants adventitious, produced at the node, along the
intemodes and just below each node, where they are sometimes paired. Leafy stems
terete, up to 2.5 mm diameter, dark green; intemodes (2-)5-20(-25) cm long. Leaves
opposite, petiolate; petiole cylindrical, slightly flattened above, 4-7 mm long, 1.5-2
mm in diameter; lamina dark green, underneath lighter green with darker midrib and
margin, fleshy, stiff (less so in cultivation), lanceolate, 4-7 x 1.5^ cm (to 10 x 5
cm in cultivation); apex acute-apiculate, base (round) acute, margin entire, midrib
and secondary veins slightly raised on adaxial surface, secondary veins 3-6 pairs,
branching at 35-60° from midrib. Inflorescences usually bearing a single open flower
at a time (up to 5 in cultivation) and 2-A developing buds; peduncle extra-axillary or
apparently axillary when borne on very short shoots, persistent, rachis one or two per
peduncle, bearing scars of previous flowerings 2-5 mm long, 2-2.5 nmi in diameter
(to 2 cm long in cultivation); pedicels 0. 6-0.8 mm long, c. 0.6 mm in diameter.
Calyx c. 2 mm in diameter, green, lobes (round) ovate 0.6-0. 8 x 0.5-0. 8 mm, apex
round, sparsely ciliate. Calycine colleter 1 or 2 at each calyx lobe sinus, ovate, with a
round apex, 100-120 x 80-100 pm. Corolla succulent, urceolate, 3.5^ x 2-2.5 mm,
basally lighter yellow, progressively fading into light pink or white at the lobe apices,
externally glabrous, corolla throat with one ring of retrorse hairs; lobes triangular,
0.5-0. 7 X c. 0.6 mm, apex acute. Corolline corona absent. Staminal corona lobes
1.5-1. 7 mm high, c. 1.5 mm in diameter, composed of five light yellow appendages
shaped like an inverted anchor with a stipe and lunate apex. Style head conical, hidden
by the anther appendages. Pollinarium erect, 550-600 x 500-600 pm with two oblong

Dischidia acutifolia a new record for Singapore

31

Fig. 1. Dischidia acutifolia in Singapore. A. The original plant as eollected by Lee and Mishak
in Sungei Buloh Wetland Reserve in 2013. B. First flowering in cultivation at SBG. C. Dissected
flower. The single ring of hairs lining the corolla throat, the yellow corona appendages and the
anther appendages covering the stigma head can be observed. D. Inflorescence with five open
flowers; usually in wild-collected plants the inflorescences have only one open flower at a
time. E. Pollinarium with twin pollinia. B-E Based on living plants cultivated at the Singapore
Botanic Gardens (SBG ace. no. 20132480). (Photos: M. Rodda)

32

Gard. Bull. Singapore 67(1) 2015

pollinia, 230-250 x 90-100 pm, apex and base rounded to obtuse, retinaculum 170-
190 X 60-70 pm, caudicles elongated, laterally crested, 250-270 pm long. Ovaty bi-
carpellate, bottle-shaped, 1-1.5 mm long, each carpel c. 0.7 mm in basal diameter.
Fruits and seeds not seen.

Singapore specimens examined. Sungei Buloh Wetland Reserve, Lee, D. & Shunari, M. s.n.,
eultivated at the Singapore Botanie Gardens, vouehered on 30 Get 2014 as Rodda, M. MR898
(SING, K, KEP).

Notes. The description of Dischidia acutifolia by Hooker (1883) was based on the
collection Maingay {Kew Distrib. 1122). The herbarium of a particular specimen
was not indicated. In Kew there is a specimen labelled as Kew distribution number
1122 which is also labelled with the Maingay numbering series (Maingay 1960). An
exhaustive search for duplicates has not been earned out but since the K specimen is
not labelled as a unicate it is possible that duplicates may be found in other herbaria
holding Maingay materials (B, BM, CAL, CGE-B, G, L [Steenis-Kruseman, 1950]).
Therefore a lectotype needs to be selected. The K specimen is well preserved and
matches the description of Dischidia acutifolia and is therefore selected as lectotype
for the name.

ACKNOWLEDGEMENTS. We thank National Parks Board, Singapore, for funding our
research, and Tanya Livshultz and one anonymous referee for their constructive comments on
the manuscript.

References

Chong, K.Y., Tan, H.T.W. & Corlett, R.T. (2009). A Checklist of the Total Vascular Plant Flora
of Singapore: Native, naturalised and cultivated species. 273 p. Singapore: Raffles
Museum of Biodiversity Research, National University of Singapore.

Davison, G.W.H. (2008). The Red List categories. In: Davison, G.W.H., Ng, P.K.L. & Ho, H.C.
(eds) The Singapore Red Data Book: Threatened plants and animals of Singapore. 2nd
ed. Pp. 1-4. Singapore: Nature Society (Singapore).

Forster, P.I., Liddle, D.J. & Nicholas, A. (1996). Asclepiadaceae. In: Orchard, A.E. (exec, ed)
Flora of Australia, vol. 28 (Gentianales). Pp. 197-283. Melbourne: CSIRO.

Hooker, J.D. (1883). Dischidia. In: Hooker, J.D. (ed) The Flora of British India 4: 49-52.
London: L. Reeve & Co.

Jagtap, A.P. & Singh, N.P. (1999). Asclepiadaceae. Fascicles of Flora of India. Fascicle 24.
Calcutta: Botanical Survey of India.

Li, P.T., Gilbert, M.G. & Stevens, W.D. (1995). Asclepiadaceae. In: Wu, Z. & Raven, P.H.
(eds) Flora of China 16 (Gentianaceae through Boraginaceae). Pp. 189-270. St. Louis:
Missouri Botanic Garden.

Livshultz, T, Tran, T.B., Bounphanmy, S & Schott, D. (2005). Dischidia (Apocynaceae,
Asclepiadoideae) in Laos and Vietnam. Bliimea 50: 113-134.

Rintz, R.E. (1980). The Peninsular Malayan species of Dischidia (Asclepiadaceae). Blumea
26: 81-126.

Dischidia acutifolia a new record for Singapore

33

Rodda, M., Tran, H.D., Chiew, RT., Liew, D. & Leong-Skomickova, J. (2012). The rediscovery
of Dischidia hirsuta (Apocynaceae, Asclepiadoideae) in Singapore. Gard. Bull.
Singapore 64: 293-299.

Steenis-Kniseman, M.J. van (1950). Malaysian plant collectors and collections: Being a
cyclopaedia of botanical exploration in Malaysia and a guide to the concerned literature
up to the year 1950. Flora Malesiana, Ser. 1,1: 5-605. Jakarta: R Noordhoff
Ziegler, A.D., Terry, J.R, Oliver, G.J.H., Daniel Friess, A., Chuah, C.J., Chow, W.T.L. &
Wasson, R.J. (2014). Increasing Singapore‘s resilience to drought. Hydrolog. Process.
28: 4543-4548.

Gardens’ Bulletin Singapore 67(1): 35-37. 2015
doi: 10.3850/S238258 12 15000034

35

From Ancistrocladus to Tristaniopsis via Tetramerista - the
convoluted history of a Wallich collection and its impact

on the native flora of Singapore

LM. Turner

Research Associate, Royal Botanic Gardens Kew,
Richmond, Surrey, UK
j_trop_ecol@yahoo.co.uk

ABSTRACT. Ancistrocladus sagittatus Wall, ex Planch. (Ancistrocladaceae) is the name of a
plant species solely based on a sterile collection made by Nathaniel Wallich in Singapore in
1822. Since 1875, this name has been treated as a variety or synonym oi Tetramerista glabra
Miq. (Tetrameristaceae). However, the original material is actually referable to Tristaniopsis
(Myrtaceae), and Ancistrocladus sagittatus is here lectotypified and reported as a synonym
of Tristaniopsis merguensis (Griff) Peter G.Wilson & J.T.Waterh. for the first time. The new
determination of the Wallich collection means that the record of Tetramerista glabra in the
native flora of Singapore requires reassessment. It is concluded that Tetramerista glabra, the
genus Tetramerista and the family Tetrameristaceae should all be excluded from inventories of
the native flora of Singapore.

Keywords. Ancistrocladus, flora, Singapore, synonymy, Tetramerista, Tristaniopsis, typification

Introduction

Nathaniel Wallich (1786-1854) was a Danish surgeon who studied botany in
Copenhagen. In 1 807, he arrived at Serampore in India to work as a medical doctor.
By 1814 he had officially joined the British East India Company, still working in the
medical service, and in 1 8 1 5 he was appointed Superintendent of the Botanic Garden
in Calcutta. Wallich undertook various expeditions to collect specimens. This included
a trip from Calcutta to Penang and Singapore in 1822. Of the many collections made
by Wallich and his assistants on this trip, one from Singapore, of apparently only sterile
material, was Ancistrocladus sagittatus by Wallich. The name was included as

number 1055 in Wallich ’s monumental listing of the East India Company Herbarium
known as the Numerical List (Wallich, 1828-1849) or Wallich’s Catalogue. As with
the vast majority of plant names in the Numerical List, Ancistrocladus sagittatus was
a nomen nudum in the absence of a description or reference to a published one. An
exception to this rule was actually provided by the generic name Ancistrocladus.
This was validly published in the Numerical List as a replacement name for Wormia
Vahl, a later homonym of Wormia Rottb. (Dilleniaceae). The top set of the East India
Company collections was maintained as a closed collection, often refeiTed to as the
Wallich Herbarium, firstly at the Limiean Society of London and later, from 1913, at

36

Gard. Bull. Singapore 67(1) 2015

the Royal Botanic Gardens Kew (K-W). Other material was distributed by Wallich
under the list numbers to many different individuals, including William Hooker, and
institutions, though it should not be assumed that all specimens with the same number
represent true duplicates.

Ancistrocladus sagittatus was first validated by the French botanist Jules Emile
Planchon in a paper on Ancistrocladus (Planchon, 1849). Planchon based the species
solely on Wallich ’s collection, but did not state which specimens he had seen. Planchon
worked as assistant to William Hooker at Kew in the period 1 844-1 848 and, for several
other Ancistrocladus species in his revision, he mdicated that he had seen specimens
in Hooker’s personal herbarium. I therefore designate a specimen distributed under
the East India Company number 1055, now in the general herbarium of the Royal
Botanic Gardens Kew and formerly in Hooker’s personal herbarium, as the lectotype
of Ancistrocladus sagittatus Wall, ex Planch.

Alfred William Bennett seems to have been the first to realise thsit Ancistrocladus
sagittatus did not belong in Ancistrocladus. In the account of the Ochnaceae for the
Flora of British India, Bennett (1875) treated Ancistrocladus sagittatus as a variety
of Tetramerista glabra Miq. (Tetrameristaceae, fonnerly placed in the Oclmaceae or
Theaceae). A casual encounter with this piqued my interest because, if this reduction
were taxonomically correct, then Ancistrocladus sagittatus, the older of the two
names, should provide the correct epithet at the rank of species. The treatment of
Ancistrocladus sagittatus as a synonym of Tetramerista glabra has been repeated in
subsequent revisions of Ancistrocladus (Steenis, 1948; Gereau, 1997; Taylor et al.,
2005) and Tetramerista (Lim, 2010).

The online catalogue of the Kew Herbarium made it easy to discover that the
recent detenninations of the Wallich specimen in the general herbarium were under
the myrtaceous genus Tristaniopsis (formerly Tristania) rather than Tetramerista. The
two genera bear superficially similar foliage. However, Tetramerista lacks a clear
intramarginal nerve (Keng, 1989), which is evident in the Wallich specimens at Kew,
leading me to agree with the placement in Tristaniopsis. Following the treatment of
Tristaniopsis by Ashton (2005, 2011), I therefore comi&QX Ancistrocladus sagittatus to
be a new synonym of Tristaniopsis merguensis Griff.

Tristaniopsis merguensis (Griff.) Peter G.Wilson & J.T.Waterh., Austral. J. Bot. 30:
439 (1982). - Tristania merguensis Griff, Account Bot. Coll. Cantor 18 (1844-1845).
TYPE: Burma, Mergui, W. Griffith 235 (isotypes K [barcode nos. K000793713,
K000793713]).

Ancistrocladus sagittatus Wall. [Numer. List no. 1055 (1829), nom. nud.] ex Planch.,
Ann. Sci. Nat., Bot. ser. 3, 13: 319 (1849), synon. nov. - Tetramerista glabra Miq. var.
sagittata (Wall, ex Planch.) A. W.Benn. in Hook.f , FI. Brit. India 1: 526 (1875). TYPE:
Singapore, 1822, N. Wallich s.n. [EIC 1055] (lectotype K [barcode no. K000793725],
designated here; isolectotypes K-W [barcode nos. KOOl 110861, K001110862]).

Tristanopsis merguensis in Singapore

37

As well as dealing with the correct placement of the namQAncistrocladus sagittatus, this
finding also brings into question the occun'ence of Tetramerista glabra in Singapore.
The species has been listed in the Singapore flora (Ridley, 1900; Keng, 1990) based
solely on the Wallich record which we now know to be an error of identification. The
work of Comer (1978) on the swamp forests of Johore and Singapore supports the
absence of Tetramerista glabra from Singapore. He found the species in the tme peat
swamp relict at Pontian in Johore, but not in the freshwater swamps in the Sedili forests
of Johore or Singapore. 1 therefore consider that there is no record of Tetramerista
glabra from Singapore and the species, genus and family all have to be removed from
listings of the native flora until evidence to the contrary is forthcoming.

References

Ashton, P.S. (2005). New Tristaniopsis Peter G. Wilson & J.T.Waterh. (Myrtaceae) from
Borneo. Gard. Bull. Singapore 57: 269-278.

Ashton, PS. (2011). Myrtaceae. In: Soepadmo, E., Saw, L.G., Chung, R.C.K. & Kiew, R.
(eds) Tree Flora of Sabah and Sarawak 7: 87-330. Kepong: Forest Research Institute
Malaysia.

Bennett, A.W. (1875). Ochnaceae. In: Hooker, J.D. (ed) Flora of British India 1: 523-527.
London: L. Reeve & Co.

Comer, E.J.H. (1978). The freshwater swamp-forest of South Johore and Singapore. Gard.
Bull. Singapore, Suppl. I: 1-266.

Gereau, R.E. (1997). Typification of names in Ancistrocladus Wallich (Ancistrocladaceae).
Novon 7: 242-245.

Keng, H. (1989). Tetrameristaceae. In: Ng, F.S.P. (ed) Tree Flora of Malaya 4: 470^71.
Petaling Jaya: Longman Malaysia Sdn. Bhd.

Keng, H. (1990). The Concise Flora of Singapore: Gymnosperms and Dicotyledons. Singapore:
Singapore University Press.

Lim, C.L. (2010). Tetrameristaceae. In: Kiew, R., Chung, R.C.K., Saw, L.G., Soepadmo, E. &
Boyce, PC. (eds) Flora of Peninsular Malaysia, Series II Seed Plants (Malayan Forest
Records 49) 1 : 265-269. Kepong: Forest Research Institute Malaysia.

Planchon, J.-E. (1849). Essai monographique d’une nouvelle famille de plantes propose sous le
nom d’Ancistrocladees. Ann. Sci, Nat., Bot. sen 3, 13: 316-320.

Ridley, H.N. (1900). The flora of Singapore. J. Straits Branch Roy. Asiat. Soc. 33: 27-196.
Steenis, C.G.G.J. van (1948). Ancistrocladaceae. In: Steenis, C.G.G.J. van (ed) Flora
Malesiana, Series 1, 4: 8-10. Jakarta: Noordhoff-Kolff N.V.

Taylor, C.M., Gereau, R.E. & Walters, G.M. (2005). Revision of Ancistrocladus Wall.

(Ancistrocladaceae). Missouri Bot. Gard. 93: 360-399.

Wallich, N. (1828-1849). A Numerical List of Dried Specimens of Plants in the East India
Company’s Museum, Collected Under the Superintendence of Dr. Wallich of the
Company’s Botanic Garden at Calcutta. London: Honourable East India Company.

Gardens’ Bulletin Singapore 67(1): 39-43. 2015
doi: 10.3850/S238258 12 15000046

39

The clarification and synonymisation of two taxa of
Vittaria from Peninsular Malaysia and a new combination
in Haplopteris (Pteridaceae subfam. Vittarioideae)

S. Lindsay^ & D J. Middleton^

'Gardens by the Bay, 18 Marina Gardens Drive,
Singapore 018953

stuart.lindsay@gardensbythebay.com.sg
^Herbarium, Singapore Botanic Gardens, National Parks Board,
1 Cluny Road, Singapore 259569
david_middleton@nparks .gov. sg

ABSTRACT. A variety and a species of Vittaria in Peninsular Malaysia are synonymised and
the new combination Haplopteris sessilifrons (Miyamoto & H.Ohba) S. Linds, is made.

Keywords. Adiantaceae, Haplopteris sessilifrons, Vittaria ensiformis var. latifolia, Vittaria
sessilifrons, Vittariaceae

Introduction

As research towards an account of Adiantaceae for the Flora of Peninsular Malaysia
is progressing, papers are being published to clarify the correct generic placement of
a number of species (Lindsay, 2010; Lindsay & Chen, 2014). Wider discussions on
the current delimitation of the Asian genera formerly placed in the family Vittariaceae
(most often treated as part of Pteridaceae subfam. Vittarioideae but to be included in
Adiantaceae for the Flora of Peninsular Malaysia) can be found in Crane (1998),
Lindsay (2004), Ruhfel et al. (2008) and Lindsay & Chen (2014).

This current paper concerns a species described by Miyamoto & Ohba (1992),
Vittaria sessilifrons Miyamoto & H.Ohba, and a variety written about by Holttum
(1955), Vittaria ensiformis var. latifolia,

Miyamoto & Ohba (1992) described Vittaria sessilifrons based on several of
their own collections from Pahang in Peninsular Malaysia. We had been unable to
locate any of the isotypes or paratypes, said to be in a number of herbaria, until it
was discovered that none of the type material had been distributed. This has now
been rectified and SING herbarium received an isotype and two paratypes in 2014.
Examination of this material has revealed it to be the same taxon previously written
about as Vittaria ensiformis var. latifolia by Holttum (1955). Unfortunately, Holttum’s
variety was not validly published due to the lack of a Latin diagnosis or description
(Art. 39.1; McNeill et al. 2012).

We had earlier delayed making a decision on what to do with the material
referable to Vittaria ensiformis var. latifolia, due to its apparent intermediate position

40

Gard. Bull. Singapore 67(1) 2015

between Haplopteris ensiformis (Sw.) E.H.Crane and H. elongata (Sw.) E.H.Crane
(formerly Vittaria ensiformis Sw. and V. elongata Sw.). Until the additional three
specimens were received from Japan, we had seen only five specimens that could
be referred to this taxon (three annotated by Holttum plus two others: Mohd Shah &
Samsuri MS3822 (SING) and R. Kiew RK1989 (KEP)). We remained unsure as to
whether it blun'ed the distinction between Haplopteris ensiformis and H. elongata,
whether it should be described as a variety of one or the other (but leaning towards H.
elongata), or whether it should be described as a new species (before we discovered
that it was already described as Vittaria sessilifrons). Holttum (1955) made it a variety
of Vittaria ensiformis, based on its sessile fronds, but noted its similarities to V.
elongata. Miyamoto & Ohba (1992), however, compared their new species to Vittaria
elongata and V scolopendrina (Bory) Sclikuhr ex Thwaites & Hook, (now Haplopteris
scolopendrina (Bory) C.Presl) although noted it ‘differs greatly’ from the latter.

With the availability of more material, now covering a broader geographical
distribution in Peninsular Malaysia, it has become clear that the characters which
define this taxon are quite stable and that Miyamoto & Ohba (1992) were quite correct
to describe it at the rank of species. It is similar to Haplopteris ensiformis in having
sessile fronds and sori in deep marginal or submarginal grooves but differs in having
a distinct costa above (at least in the lower half of the frond) and a wider and thinner
lamina in which the lateral veins are visible (at least with transmitted light). It is similar
to Haplopteris elongata in the width and texture of the lamina, the visibility of the
venation and the position and structure of the sori, but differs in having sessile fronds
and costae that are more strongly raised above (the costae of H. elongata are usually
distinct but flat or hardly raised above). It is similar to Haplopteris scolopendrina in
the texture of the lamina and the visibility of the venation but differs in having much
narrower and shorter fronds and sori in deep marginal or submarginal grooves (the sori
of H. scolopendrina are in shallow and broad submarginal grooves).

Based on the arguments to be found in Crane (1998) and Lindsay (2004), Vittaria
sessilifrons now requires a combination in Haplopteris which is here provided.

New combination

Haplopteris sessilifrons (Miyamoto & H.Ohba) S.Linds., comb. nov. - Vittaria
sessilifrons Miyamoto & H.Ohba, Acta Phytotax, Geobot. 43(1): 33, f. 2 (1992). -
TYPE: Peninsular Malaysia, Pahang, Gunung Tahan, Sungai Luis, alt. 700 m, 19
March 1990, Ohba, H. d Miyamoto, F 900536 (holotype Tl!; isotypes FRI, L, US,
SING!).

Vittaria ensiformis var. latifolia Holttum, Rev. FI. Malaya 2: 614 (1955 [‘1954’]),
nom. inval.

Epiphytic or lithopyhtic. Rhizome short-creeping, usually obscured by a mass of
extremely hairy roots, c. 1-2 mm diameter (in the dry state), bearing fronds close

New combination in Haplopteris

41

together, densely scaly. Scales linear, stiff, straight, gradually narrowing from a
cordate base towards a long-tailed and ultimately filifonn apex, up to c. 6 mm long and
c. 0.7 mm wide (at base), clathrate, sparsely toothed at margin particularly near base
and somewhat bicoloured, the walls of the inner cells being thicker and darker than
those of the outer cells (most noticeable in the section of the tail that is two cells wide;
the common wall in the middle is thick and black while the two marginal walls are
thinner and somewhat red). Fronds pendulous, simple, linear, 9^0 cm long, 0.45-1 .3
cm wide, gradually nan*owing towards both ends, the apex narrowly acute, the base
sessile. Lamina coriaceous, glabrous, the margins revolute almost the entire length
of the frond, midrib and margins moderately to strongly raised on the upper surface
particularly in the lower half of the frond, the midrib usually becoming flatter towards
the frond apex but the margins remaining raised throughout, midrib flat and indistinct
below. Lateral veins more-or-less visible from above (or with transmitted light),
indistinct below, simple, very oblique, parallel, joined by a continuous submarginal
vein. Sori very long, linear, uninterrupted, appearing to be marginal (but actually arising
from the submarginal vein), in a deep, two-lipped, groove in the revolute frond margin
(therefore only visible from below), the two lips more-or-less equal. Sporangia with
annuli composed of 16-18 cells. Soral paraphyses copious, their stalks multicellular,
filiform, mostly colourless; their apical cells large, funnelform, twice as long as wide
or longer, reddish-brown. Stint ctures resembling sporangial stalks (but without
sporangia attached) more numerous than paraphyses. Spores bilateral, bean-shaped,
monolete, pale, translucent, smooth.

Distribution. Peninsular Malaysia and, possibly, Borneo. We have seen specimens from
Johor, Negeri Sembilan, Pahang and Terengganu (close to the border with Kelantan)
in Peninsular Malaysia. Holttum (1955) lists Perak too. We have not verified material
from Borneo listed as Vittaria ensiformis var. latifolia in online herbarium specimen
databases.

Ecology. Information is scant. However, three collections are described as epiphytic
on mossy tree-trunks near streams in forests, one as epiphytic 3 m from the ground
above a river, and one as growing on rocks. The lowest recorded altitude is 200 m and
the highest is 762 m.

Provisional lUCN consen^ation assessment. Data Deficient (DD). Although not often
collected, this species is widespread in Peninsular Malaysia which would suggest an
assessment of Least Concern. However, of the collections known, several are more
than 100 years old and none have been collected since 1990.

Additional specimens examined. PENINSULAR MALAYSIA: Johor: Kota Tinggi, [locality
illegible], 1910 [month illegible], Ridley, H.N. s.n. (SING [SING0033264]); Mount Austin,
[1901?; date illegible], Ridley, H.N. s.n. (SING [SlNG008566i]); Sungai Salat, Ulu Endau,
Sep 1985, Kiew, R. RKJ989 (KEP). Negeri Sembilan: Gunung Angsi, alt., 2500 ft. [762 m], 22
Nov 1923., M/r. Md SFN11617 (SING). Pahang: Gunung Tahan, Sungai Juram, alt. 200 m, 11

42

Gard. Bull. Singapore 67(1) 2015

Mar 1990, Ohba, H. & Miyamoto, F 900120 (SING, TI); Smigai Juram to Sungai Luis, alt. 700
m, 13 Mar 1990, Ohba, H. & Miyamoto, F. 900208 (SING, Tl). Terengganu: Gunung Ayam
via Ulu Besut, alt. 2,400 ft. [732 m], 5 Mar 1976, Shah, Mohd & Samsuri MS3822 (SING).

Notes. The holotype and SING isotype are clearly labelled as having been collected on
19 March 1990; the date in the protologue of 13 March 1990 is a printing error.

In his key to the Malayan species of Vittaria, Holttum (1955: 608) says of
Vittaria ensiformis var. latifolia that the “midrib [is] never strongly raised on [the]
upper surface”. This statement is at odds with the protologue of Vittaria sessilifrons
which says “costa and margin raised on upper side” and illustrates both as being rather
strongly raised (see Miyamoto & Ohba, 1992, Fig. 2E & G). We have examined three
specimens annotated by Holttum and, in each, the costa and margins are moderately to
strongly raised on the upper surface of the basal half of the frond.

Miyamoto & Ohba (1992) state that Vittaria sessilifrons differs greatly from V
scolopendrina in having a raised costa. This implies that Flaplopteris scolopendrina,
using the cuiTent name, does not have a raised costa when in reality it does. The costa
of Haplopteris scolopendrina is, in fact, very strongly raised on the upper surface of
the frond (even more so than in Flaplopteris sessilifrons) for almost the entire length
of the frond.

The illustration of the paraphyses in the protologue of Vittaria sessilifrons shows
them with ovoid or clavate heads but we have not seen any paraphyses of this sort in
the type material or in any other material. Instead, all material that we have examined
has paraphyses with funnelform apical cells, a shape that is typical for Haplopteris. As
well as these paraphyses, all of the specimens of Haplopteris sessilifrons have a very
large number of structures that look like sporangial stalks without sporangia attached.
It is unclear whether these are an artefact of dried specimens or if, as we suspect, these
develop without ever producing sporangia. Similar structures are sometimes found in
both Haplopteris elongata and H. ensiformis but never so densely.

ACKNOWLEDGEMENTS. We thank Tetsuo Ohi-Toma (TI), Akiko Shimizu (TI), Futoshi
Miyamoto (Tokyo University of Agriculture) and Serena Lee (SING) for arranging the transfer
of type material from TI to SING. We also thank Jana Leong-Skomickova for her valuable
comments and for guiding this paper through the review process.

References

Crane, E.H. (1998 [‘1997’]). A revised circumscription of the genera of the fern family
Vittariaceae. Syst. Bot. 22(3): 509-517.

Holttum, R.E. (1955 [‘1954’]). A Revised Flora of Malaya, edition 1, vol. 2 Ferns of Malaya.
Government Printing Office, Singapore.

Lindsay, S. (2004). Considerations for a revision of the fern family Vittariaceae for Flora
Malesiana. Telopea 10(1): 99-112.

New combination in Haplopteris

43

Lindsay, S. (2010). New combinations m Haplopteris (Adiantaceae) for the Flora of Peninsular
Malaysia. Gard. Bidl. Singapore 62(1): 119-120.

Lindsay, S. & Chen, C.W. (2014). Three new combinations m Haplopteris (Pteridaceae subfam.
Vittarioideae). Gard. Bull. Singapore 66(2): 169-171.

McNeill, J., Barrie, F.R., Buck, W.R., Demoulin, V., Greuter, W., Flawksworth, D.L., Herendeen,
P.S., Knapp, S., Marhold, K., Prado, J., Prud’homme van Reine, W.F., Smith, G.F.,
Wiersema, J.H. & Turland, N.J. (2012). International Code of Nomenclature for algae,
fungi and plants (Melbourne Code). Regnum Vegetabile 154. 205 pp. Konigstein: Koeltz
Scientific Books.

Miyamoto, F. & Ohba, H. (1992). Two new ferns from Gunung Tahan, Malay Peninsula. Acta
Phytotax. Geobot. 43(1): 31-35.

Ruhfel, B., Lindsay, S. & Davis, C.C. (2008). Phylogenetic placement of Rheopteris and the
polyphyly of Monogramma (Pteridaceae s.l.): Evidence from rbcL sequence data. Syst.
Bot. 33(1): 37^3.

Gardens’ Bulletin Singapore 67(1): 45-49. 2015
doi: 10.3850/S2382581215000058

45

Bothriospermum (Boraginaceae), a new generic record for

the Flora of Peninsular Malaysia

A.R. Rafidah, S. Syahida-Emiza, R. Kiew & K. Imin

Forest Biodiversity Division, Forest Research Institute Malaysia (FRIM),

52109 Kepong, Selangor, Malaysia
rafidahar@fi'im.gov.my

ABSTRACT. Bothriospermum zeylanicum (J.Jacq.) Druce (Boraginaceae), a species native
to Asia, has recently become naturalised in Cameron Highlands, Pahang, Malaysia. A
description of the genus and species together with colour photographs are provided. It is the
only representative of Boraginaceae sensu stricto (i.e. subfam. Boraginoideae) in Peninsular
Malaysia.

Keywords. Boraginaceae, Bothriospermum, naturalised weed, new genus record, taxonomy,
Malaysia

Introduction

In 2007 a fieldtrip to Cameron Highlands, Pahang, Penmsular Malaysia led to the
discovery of a species of Boraginaceae that was tentatively identified as a Myosotis, a
genus not otherwise recorded from Peninsular Malaysia. Realising it was something
new, a second fieldtrip was organised in April 2014 to obtain more material and to take
photographs so that it could be correctly identified. It proved to be Bothriospermum
zeylanicum, which is a new record for the genus and species for Peninsular Malaysia;
the genus is not recorded in either the Flora Malesiana revision of the Boraginaceae
(Riedl, 1997), nor in the checklist by Turner (1997). New weeds, often of temperate
species, are regularly introduced into the Cameron Highlands, where extensive
vegetable and flower farms have replaced the lower montane forest. For example,
Kiew (2009) recently reported 17 new records of naturalised weed species, of which
ten were from Cameron Highlands. Interestingly, Bothriospermum zeylanicum occurs
at a lower altitude (c. 1200 m) than most of these other weeds. A ftdl morphological
description of the genus and species, together with colour photographs, is provided
here.

Turner (1997), in his catalogue of Peninsular Malaysian plants, listed under
Boraginaceae 16 species in 9 genera: Argusia (1 species), Carmona (1 species),
Coldenia (1 species), Cordia (4 species), Ehretia (4 species), Heliotropium (1
species), Pteleocarpa (1 species), Rotula (1 species) and Tournefortia (2 species).
Recent molecular work by Refulio-Rodriguez & Olmstead (2014) separated Cordia
and Coldenia into Cordiaceae; Ehretia, which now includes Carmona and Rotula
(Gottschling et ah, 2014), into Ehretiaceae; 'md Argusia, Heliotropium and Tournefortia

46

Gard. Bull. Singapore 67(1) 2015

into Heliotropiaceae. Pteleocarpa is now placed in Gelsemiaceae (Struwe et al.,
2014). In other words, all the genera listed by Turner under Boraginaceae have been
assigned to other families meaning that now the Boraginaceae s.s. is represented in
Malaysia only by this new record, Bothriospermum zeylanicum. Even if the narrow
family circumscription adopted by Refulio-Rodriguez & Olmstead (2014) is not
followed and one prefers to recognise Cordiaceae, Ehretiaceae and Heliotropiaceae
as subfamilies Cordioideae, Ehretioideae and Heliotropoideae (e.g. Engler & Prantl,
1897; APG III, 2009), it remains true that Bothriospermum zeylanicum is the only
Malaysian representative of the type subfamily Boragmoideae.

Taxonomic treatment

Bothriospermum Bunge, Enum. PI. China 47 (1833); Clarke, FI. Brit. India 4: 167
(1885); Backer & Bakhuizen /, FI. Java 2: 463 (1965); Zhu et al., FI. China 16: 418
(1995); Riedl, FI. Malesiana ser. I, Spemiat. 13: 63 (1997); Hsiao & Liu, FI. Taiwan 4:
387 (1998). - TYPE: Bothriospermum chinense Bunge.

Annual or biennial herbs, hispid or pubescent. Stems erect or prostrate. Leaves spirally
arranged; lamina elliptic, ovate or oblanceolate, margin entire. Flowers extra-axillary,
small, solitaiy but often fonning a leafy raceme in distal part of the branches, distinctly
pedicellate; calyx 5-lobed, the lobes divided to the base, narrowly lanceolate, slightly
enlarged in fruit; corolla blue or white, rotate, tube short, lobes 5, spreading, obtuse,
5 small scale-like appendages present in the throat. Stamens included in the corolla,
filaments very short, anthers 5, ovoid. Ovary deeply 4-lobed, style short, persistent,
stigma small, capitate; gynobase flat. Fruits-, nutlets 4, veiy small, erect, ellipsoid or
subglobose attached to the flat or nearly flat receptacle, surface usually sculptured,
ventral side with a large aperture surrounded by prominent margin.

Distribution. About 5 species, distributed in Afghanistan, Kazakhstan, Kyrgyzstan,
Tajikistan, Turkmenistan, Uzbekistan, Russia, Pakistan, India, China (its centre
of distribution), Japan, Korea, Taiwan, Vietnam and the Philippines; naturalised in
Indonesia (Java).

Bothriospermum zeylanicum (J.Jacq.) Druce, Bot. Exch. Club Brit. Isles 4: 610
(1917); Zhu et al., FI. China 16: 419 (1995); Riedl, FI. ser. I, Speraiat. 13: 63 (1997);
Hsiao & Liu, FI. Taiwan 4: 388 (1998). -Anchusa zeylanica J.Jacq., Eel. PI. Rar. 1(3):
47, t. 29 (1812). - TYPE: Plate t. 29 in Jacquin, Eel. PI. Rar. 1(3) (1812). (Fig. 1 & 2)

Anchusa tenella Homem., Hort. Bot. Hafii. 1: 176 - Bothriospermum tenellum

(Homem.) Fisch. & C.A.Mey., Index Sem. Hort. Petrop. I: 23 (1835); Clarke, FI. Brit.
India 4: 168 (1885); Backer & Bakliuizen, f, FI. Java 2: 463 (1965). - TYPE: Herb.
Vahl s.n. ‘Hab. in China’ (holotype C n.v.).

Bothriospermum, a new generic record for Peninsular Malaysia

47

Fig. 1. Bothriospermum zeylanicum (JJacq.) Druce. A-B. Habit. C. Arrangement of flowers
and fruits. D. Nutlet. E. Leaf F. Flower. Scale bars: A, B, C (5 cm); D (1 mm); E (2 mm); F (5
mm). (Photos: A.R. Rafidah)

48

Gard. Bull. Singapore 67(1) 2015

Fig. 2. Habitat of Bothriospermum zeylanicum (JJacq.) Druce. (Photo: A.R. Rafidah)

Short-lived herbs with prostrate stems. Stems slender, densely pubescent throughout,
much branched, up to 50 cm long. Leaves: petiole short, c. 5 mm long in lower leaves,
lacking in the upper; lowermost laminas 20-30 x 6-10 mm, decreasing in size to 4-10
X 2-4 mm in the upper ones, elliptic to lanceolate, margin slightly undulate, densely
strigose on both surfaces, base cuneate, midrib impressed above, prominent beneath,
apex acute. Pedicels c. 2 mm long. Flowers: calyx green, densely hairy outside;
corolla white or very pale purple (in Peninsular Malaysia), c. 2 mm long, lobes 5,
longer than the tube, c. 2 mm long, apex broadly rounded, throat appendages white,
trapeziform, emarginate, c. 0.2 mm. Stamens 5, included, inserted at the middle of
corolla tube, filaments short, anther yellowish turning brown. Ovary green, style less
than 1 mm long, terete. Nutlets green turning brown, ellipsoid, to 1 mm long, aperture
longitudinally elliptic, surface warty; calyx persistent, c. 3 mm long.

Distribution. As for the genus, in Peninsular Malaysia only from Cameron Highlands,
Pahang.

Ecology. Growing in waste land on sandy soil or in open fields or near rivers, at
c. 1200 m altitude.

Bothriospermum, a new generic record for Peninsular Malaysia

49

Specimens examined. PENINSULAR MALAYSIA: Pahang: Cameron Highlands, Belati
Estate, 4°27'N 101°28'E, 8 Oct 2007, Kiew et al, FRI70433 (KEP), Kampung Terla, 4°31'N
I01°23'E, 8 Oct 2007, Kiew et al, FRI70473 (KEP), Habu, 4°26'N I0I°23'E, 23 Apr 2014,
Rqfidah etal, FR175968 (KEP), Habu, 4°26'N I01°23'E, 23 A^x2(dU,Rafidah etal, FRI75969
(KEP).

ACKNOWLEDGEMENTS. This research was carried out as part of the Documentation and
Inventory Flora Malaysia project (Flora of Peninsular Malaysia) funded by the Ministry of
Science, Teclmology and Innovation at the Forest Research Institute Malaysia (FRIM), Kepong.
We are grateful to the curators and staff of the herbaria at BO, K and KEP for permission to
examine specimens in their care and for providing information and images. We thank Dr Saw
Leng Guan and Dr Richard Chung for their helpful comments and two anonymous reviewers
for their valuable comments on the manuscript.

References

APG III. (2009). An update of the Angiospenn Phylogeny Group classification for the orders
and families of flowering plants: APGIII. Bot. J. Linn. Soc. 161: 105-121.

Engler, H.G.A, & Prantl, K. (1897). Die NatiirJichen Pflanzenfamilien. Leipzig: W. Engelmann.
Gottschling, M., Luebert, F., Hilger, H.H. & Miller, J.S. (2014). Molecular delimitations in the
Ehretiaceae ( Boraginales). Mo/^c. Phylogenet. Evol. 72: 1-6.

Kiew, R. (2009). Additions to the weed flora of Peninsular Malaysia. Malayan Nat. J. 61:
133-142.

Refulio-Rodriguez, N.F. & Olmstead, R.G. (2014). Phylogeny of Lamiidae. Amer. J. Bot.
101(2): 287-299.

Riedl, H. (1997). Boraginaceae. Flora Malesiana, Ser. 1, 13; 43-144.

Struwe, L., Soza, V.L, Sugumaran, M. & Olmstead, R.G. (2014). Gelsemiaceae (Gentianales)
expanded to include tire enigmatic Asian genus Pteleocarpa. Bot. J. Linn. Soc. 175:
482-496.

Turner, I.M. (1997 [‘1995’]). A catalogue of the vascular plants of Malaya. Card. Bull.
Singapore 41: 1-757.

Gardens’ Bulletin Singapore 67(1): 51-60. 2015
doi: 10.3850/S238258121500006X

51

Novitates Bruneienses, 1. A background to the botanical
survey of Brunei Darussalam, and a new species of

Jarandersonia (Malvaceae)

A. A. Joffre^, A.K. Muhammad Ariffin^, Y.W. Low^ & K.M. Wong^

^Brunei National Herbarium, Forestry Department,

Ministry of Industry and Primary Resources, Jalan Menteri Besar,
Berakas, BB3910 Brunei Darussalam
^Herbarium, Singapore Botanic Gardens, National Parks Board,

1 Cluny Road, 259569 Singapore
wkm2000@gmail.com

ABSTRACT. A brief introduction to research and botanical documentation of the Brunei
flora, and the collaborative programme for a continuing botanical survey of the country, is
given. An outline of the key biogeographical features of the Brunei area supports the premise
that distinct geo-ecological enclaves occur which are special units of species richness, within
which a signiflcant level of undocumented plant diversity still exists. Jarandersonia yahyantha
K.M. Wong, Joffre, Ariffin & Y.W.Low (Malvaceae), a new tree species so far only known in
Brunei, is described.

Keywords. Biological hotspot, biogeography, Borneo, Malesia, new species. Southeast Asia

Introduction: The floristic richness of Borneo and Brunei Darussalam

This new series supports a recently established programme to further the botanical
exploration of Brunei Darussalam, in northwestern Borneo, the third-largest island in
the world and biologically the richest landmass of the Sundaland region (MacKinnon
& MacKinnon, 1986; Ashton, 1989). The Malesian region as a whole (Malay
Peninsula and Malay Archipelago, including the area from the Philippines through to
New Guinea) has a diverse flora estimated to include 42,000 seed plant species (Roos,
1993). Myers et al. (2000) considered Sundaland, which includes the Malay Peninsula,
Sumatra, Java and Borneo, one of tropical Southeast Asia’s four biodiversity hotspots.

Estimates of the Bornean vascular flora have ranged from 9000 (Merrill, 1921)
to 15,000 species (Merrill, 1950; Raes et al., 2009). However, although recent work
completed by Beaman & Anderson (2004) enumerated more than 5000 vascular plant
species in Kinabalu Park (Sabah) alone, further work has suggested that many more
novelties were to be expected there (Chen et al., 2014), as well as generally with
the Tree Flora of Sabah and Sarawak (e.g., Soepadmo & Wong, 1995 and further
volumes). As such, Merrill’s upper limit for Bomeo, or a tally in excess of that, is to
be expected. On the other hand, Brunei, with its land area of 5765 knP being less than
a mere 1% of the whole of Bomeo, was estimated to have close to 5000 species of

52

Gard. Bull. Singapore 67(1) 2015

seed plants, including 2000 species of tree (Wong, 1997). A first intensive checklisting
of the flora based on organised field collecting over some seven years, 1989—1995,
much of it an active collaboration between the Brunei Forestry Department and the
Royal Botanic Gardens, Kew (Coode et ah, 1996), accounted for some 3500 species
of indigenous seed plants, including 1900 tree species (Wong, 1999). Apart from the
Dipterocarpaceae, which has been well treated by Ashton (1964, 1 982) and is generally
well collected, this checklist remains a first-stage investigation of the flora because (1)
many families and genera still had significant numbers of undetennined specimens;
(2) recent critical revisions were not available then for many genera; and (3) in the
ensuing period the Brunei Herbarium continued to gather interesting taxa that had not
been captured by earlier collecting effort. Thus it can be appreciated that the Brunei
flora is a very rich one.

In 2013, the Government of His Majesty The Sultan of Brunei Damssalam
and the National Parks Board, Singapore, entered into a collaborative programme for
a new phase of botanical survey. This seeks to intensify fieldwork for biodiversity
exploration and documentation, and to focus on the botanical inventory of Brunei,
including taxonomic identifications by specialists from our own and outside institutions
when possible.

Key biogeographical features of the Brunei area

In the same year as this new collaboration began, a review of the flowering plants
endemic to Brunei was published (Henrot et al., 20 1 3), listing 65 species of angiospemis
hitherto known only from Brunei, and suggesting that, as with a number of other species
generally occurring only in the Brunei area (which includes adjacent parts of SW Sabah
or NE Sarawak), some of the so-called endemics could subsequently be found to be
not so restricted. The latter category is exemplified by the aroids Homalomena scutata
S.Y.Wong & P.C. Boyce, occuning in both the Belait and Temburong districts of
Brunei but found also in the Limbang and Mulu areas in adjacent NE Sarawak (Wong
& Boyce, 2014), and H. cowleyae RC, Boyce & S.Y.Wong, known in the Temburong
and Mulu areas (Boyce & Wong, 2014),

There are at least two biogeographic scenarios why this work on endemic
plants is significant. One is that we could indeed expect there would be true Brunei
endemics, as botanical experience has shown for, especially, herb species, including
many Begonia or gesneriad taxa, as well as smaller plants in general. Henrot et al.
(2013) found that most of the documented endemic taxa were smaller plants in the
Begoniaceae (15 taxa), Araceae (8 taxa), Gesneriaceae (7 taxa), Orchidaceae (5 taxa),
Zingiberaceae (5 taxa), and palms (Arecaceae: 5 taxa), so it could be true that many
large flowering plants have better dispersal than small ones. Although it should be
borne in mind that, thus far, the collecting and research effort has concentrated much
on trees, rather than herbs, other reasons, including recent evolution, may restrict
plant distribution. A slender bamboo, Temhurongia simplex S.Dransf. & K.M.Wong,
is the only known representative of this peculiar genus (Dransfield & Wong, 1996)

A new Jarandersonia from Brunei Darussalam

53

that is still entirely restricted to Brunei’s Temburong valley, even after two decades
of continuing botanical collecting in Borneo. There is a suite of similarly restricted
Begonia and other species. Some others are borderline cases: for instance, the palm
Livistona exigua Dransf, documented from Bmnei’s Batu Patam ridge, still has not
been documented from adjacent parts in Sarawak.

Wong (1997, 1 999) discussed how the Brunei area includes its own geographical
and ecological enclaves within the geologically distinct (but relatively youthful) and
larger northwest Borneo area, taking into consideration how its main area is hemmed
in by either highlands (such as around inland Temburong, Tutong and Belait) or
extensive swamp systems (as in the lower Belait and Baram drainage complex to its
southwest) (Fig. 1). The western side of Brunei is largely the remains of an ancient
syncline, now drained by the Belait and Tutong rivers, whereas the eastern Temburong
district is mostly the drainage area of one major river system, the Temburong (Ashton,
1958). This, together with precipitous topogi*aphy marked by steeply incised valleys
in the hard sandstones of inner Temburong, or in the uppermost reaches of the Belait
and Tutong, can conceivably bring about population isolation that engenders highly
localised endemism (Coyne & Orr, 2004). Even in the Belait district, where topographic
relief is generally lower than in the Temburong, hilly pockets isolated by swamps or
river systems, or highly specific habitats such as moist sandstone bluffs, may harbour
narrowly endemic or habitat-specialist taxa, such as Homalomena spp. documented
only from the Teraja area of Belait (Wong & Boyce, 2011). Many of the flowering
plants endemic to Brunei Darussalam are either very rare or undercollected: 15 are
known from a single collection only and 83% are known from three sites or fewer.

The other significant dimension is that Bmnei has maintained maximal levels
of forest cover within the country (Bryan et ah, 2013). Borneo’s forest cover declined
from 71% of the island (mid-1980s) to 54% by 2000, an alarming rate of deforestation
(Stibig & Malingreau, 2003). With continuing rainforest degradation in the region,
Brunei Darussalam could continue to serve as a critical refuge for Bornean plants
impacted by such processes. Henrot et al. (2013) discussed the increasing importance
of ‘anthropogenic endemics’ in refuge in Brunei fi*om the wider Brunei area or beyond:
it is a scenario we would not wish to see exacerbated, but the significance of conserved
resources in Brunei becomes obvious. In upping the ante, the Brunei Ministry of
Industry and Primary Resources phased out loggmg in all Forest Reserves (previously
some had been classified for timber production) in 2014 (Brunei Times, 2014).

Continuing efforts

All this underscores how important it is now for authorities in Borneo to redouble
efforts into biodiversity inventory and conservation planning, in order that as many
specially diverse or unique areas as possible can be recognised in time and be
brought under effective management. For our part, it brings us back to the special
collaboration now in place between the Brunei National Herbarium (acronym BRUN;
Thiers, continuously updated) and the Singapore Herbarium (SING), and their partner
institutions, for continuing the Botanical Survey of Bmnei Damssalam.

54

Gard. Bull. Singapore 67(1) 2015

Fig. 1. The Brunei area: Key geological, tectonic and topographical features.

Uncoloured'. Pliocene - Miocene Sediments, in places overlain by Pleistocene or more recent
deposits. Yellow. Miocene - Oligocene Sediments. Brown: Rajang Group Deepwater Sediments
(Late Cretaceous - Eocene). Anticlines: Al. Belait anticline. A2. Jerudong anticline. A3.
Danau anticline. A4. Tangga anticline. A5. Mulu Uplift. Synclines: SI. Liku-Badas syncline.
S2. Belait syncline. S3. Limbang syncline. S4. Labu syncline. S5. Tangga syncline. Simple
line with arrows: Approximate boundary of synclinal basins within Brunei. PS. Peat body
or peatswamp vegetation over low-lying ground. Map prepared by Alsa Moo & K.M. Wong,
based on information from Liechti (1960), Wilford (1960) and Tate (2002).

In this series, we document new botanical findings in Brunei advanced through
our research or that of our collaborators: the new taxa discovered or diagnosed, the
new records of plants not previously documented in Brunei, and new and noteworthy
observations. It is our hope, given the indications from past and ongoing studies,
that the Brunei area continues to yield new and interesting plant taxa, and that such
a series will help focus due attention on Brunei’s floristic richness. As the present
paper shows, both newer (post Checklist, Coode et ah, 1996) as well as older (Ashton)
collections from Brunei are still important for our continuing studies, attesting to the
value of a good botanical archive in helping to attain further or newer biodiversity

A new Jarandersonia from Brunei Darussalam

55

documentation. All this is permitted only by sustained botanical effort and resources
over the longer time frame, by necessity trans-generational in scope.

A new species of Jarandersonia (Malvaceae)

Jarandersonia yahyantha K.M. Wong, Joffre, Ariffin & Y. W.Low, sp. nov.

This new species is most similar to Jarandersonia pannfolia Kosterm. in having fruits
with slender spines that are tuberculate and which bear tufted-hairy indumentum, but
differs in its subcordate to cordate (not cuneate to rounded) leaf bases and fruit spines
of 25-35 mm (not 10-15 mm) long. - TYPE: Brunei, Belait, Sg Liang, Andulau Forest
Reserve, Compartment 18, Labi Road, 10 April 2014, Ariffin, Jangarun & Raiizaidi
BRUN 24174 (holotype BRUN; isotypes A, K, KEP, L, SAN, SAR, SING). (Fig. 2)

Medium sized tree 10-25 m tall, trunk slightly fluted; bark smooth, pale grey brown.
Leaves broad-elliptic, 4.5-17(-22.7) cm long, 3.5-8(-11.5) cm wide; base subcordate
to cordate; apex acute to rounded to obtuse to slightly emarginate; midrib flat to slightly
sunken on upper surface, prominent on lower surface, secondary veins 6-8 pairs,
making angular loops near the leaf margin, the basal 1-2 pairs making a smaller angle
with the midrib than other secondaries (so resembling basal veins in a 3- to 5 -nerved
leaf base, but not reaching a fifth of the leaf length), flat to very slightly prominent
on upper side, prominent on lower side; glabrous on entire upper surface, with dense,
overlapping stellate scales completely occluding the lower surface; petioles 7-15 mm
long, 1.5-2 mm diameter, densely stellate lepidote. Inflorescences axillary, borne in
distal leaf axils, on young leafless imiovations or more proximal twig portions that
have lost their leaves, paniculate, to (l,5-)4-9,5 cm long, bearing (2-)4-9 flowers;
bracts ovate, acute, 2-4 mm long, 1.5-2 mm broad; flower pedicels (3-)5-9 mm long
(7-11 mm long in fresh material), scaly; calyx tube c. 2.5^ mm long and 2. 5-3. 5 mm
diameter (3^.5 mm long and 5-6 mm diameter in fresh material), scaly outside (scale
margins subentire-erose to short-fimbriate), lobes triangular, 2. 5-3. 5 mm long and
2. 5-3. 5 mm wide at base (6-7 mm long and 3-4 mm wide at base in fresh material),
scaly outside; petals 5, obovate, (5. 5-) 7-9 mm long and 3-6 nnm wide (16-18 nmi
long and 8-12 mm wide in fresh material), plane to slightly incurved, creamy white;
stamens many, filaments 3-7 mm long (7-1 5 mm long in fresh material), pale greenish
yellow, anthers knobby, c. 0.3 mm long, pale yellow; ovary subglobose, 5-lobed when
dried, 1-1.5 mm diameter (2-3 mm diameter in fresh material), pale greenish yellow,
short-hairy. Infructescence 3.5-13 cm long. Fruit subglobose, 6-10 mm diameter,
3 -locular, outside covered with slender soft spines 25-35 mm long, spines tuberculate
and bearing tufted hairs; seeds l(-2), ovoid-elipsoid, 17 mm long, 4 mm wide, dark
brown, smooth.

Distribution. The new species is apparently endemic to Brunei Darussalam. Of the
other species in the genus, only Jarandersonia parvifolia, which has a widespread
distribution in Sarawak, has been recorded in Brunei (Jaamat & Tachun FMS 39640,
KEP) in the Labi Forest Reserve.

56

Gard. Bull. Singapore 67(1) 2015

Fig. 2. Jarandersonia yahyantha K.M.Wong, Jof&e, Arififin & Y.W.Low. A. Tree in flower. B.
Flowering braneh. C. Fallen fiaiits (some germinating) and leaves. A & B from Ariffin et al. BRUN
24087; C from Ariffin et al. BRUN 24174. (Photos: A-B: Muhammad Ariffln, C: K.M. Wong)

A new Jarandersonia from Brunei Darussalam

57

Provisional lUCN conservation assessment. The provisional lUCN conservation
status for Jarandersonia yahyantha, assessed with the aid of GeoCAT (Bachman et ah,
2011), is Critically Endangered (CR B2ab(iii); D) (lUCN, 2012). The ‘B2’ designation
results from an area of occupancy (AGO) estimated to be less than 10 km^ (about 8
km2 for J. yahyantha); ‘a’ is due to a fragmented distribution as it occurs in only two
populations, namely at Bukit Jerudong and Andulau Forest Reserve; ‘b(iii)’ is due
to reductions in the area and quality of its habitat as J. yahyantha is so far known
only in the Compartments 7 and 18 of the Andulau Forest Reserve, which has some
disturbance along part of their boundary by adjacent road development, and on Bukit
Jerudong near Kampung Peninjau, which is previously disturbed forest; ‘D’ is due to
the small population size of only four mature individuals known so far.

Etymology’. The new species honours Pehin Orang Kaya Seri Utama Dato Seri Setia
Awang Haji Yahya bin Begawan Mudim Dato Paduka Haji Bakar, Brunei Darussalam’s
Minister of Industry and Primary Resources, for his interest in conservation and
leadership in ceasing all timber production from Bmnei’s forest reseiwes.

Additional specimens examined. BRUNEI. Belait: Simgai Liang, Andulau Forest Reserve,
Compartment 1 8, Labi Road, 200-300 m from roadside, 28 May 2014, Ariffin, Watu & Khairul
BRUN 24112 (A, BRUN, K, KEP, L, SAN, SAR, SING), 24112 (BRUN, K, SAN, SING).
Brunei-Muara: Jerudong, Kampung Peninjau, Bukit Jerudong, 11 Mar 2014, Ariffin, Watu,
Azlan & Khairul BRUN 24087 (BRUN, K, L, SAN, SAR, SING).

Notes. There are six known species of this genus (Tan et al., 201 1; Chung et al., 2012),
established by Kostermans (1960), in which the fruits are typically covered with
spines bearing setose hairs. Of these, the new species here is most like Jarandersonia
pentaceoides R.C.K.Chung & H.S.Tan, J. paimfolia and J. rinoreoides Kosterm. in
having slender soft fruit spines (the other species, J. clemensiae (Burret) Kostenn., J.
pursglovei (Kosterm.) Kosterm. and J. spinulosa Kosterm., have short stout and stiff
fruit spines.) The new species here differs from Jarandersonia pentaceoides in having
short basal secondary veins that do not reach even a fifth of the length of the leaf (in
that species the basal secondary veins are conspicuous in reaching halfway or more
along the leaf length) and fruit spines that are 25-35 mm long (the fruit spines in that
species are only 10-20 mm long). It differs from Jarandersonia rinoreoides in having
subcordate to cordate (not typically cuneate) leaf bases and fruit spines that are 25-35
mm (not 6-28 mm) long and short-tuberculate (rather than non-tuberculate); and from
J. parxAfolia as mentioned above in the diagnosis. Leaf size is not a particularly reliable
character because the flowering or fruiting material taken for herbarium specimens
mostly occurs on higher branches bearing smaller leaves, although larger leaf blades
may also be occasionally found on such branches.

Note that Tan et al. (201 1 ) and Chung et al. (2012) refer to the tuberculate spines
as “short-branched”. Also, the regularly polygonal fine vein areoles on the lower leaf
surface of Jarandersonia pentaceoides, which they described as “distinct honey comb-
like quaternary veins”, is not restricted to that species, being also found on occasion
in J. rinoreoides (e.g., Dewol SAN 99462 in SING); this feature is, however, absent

58

Gard. Bull. Singapore 67(1) 2015

from the leaves of the new species described here, in which the lower leaf surface is
completely occluded by dense, overlapping scales.

ACKNOWLEDGEMENTS. We would like to express our gratitude to Pehin Orang Kaya Seri
Utama Dato Seri Setia Awang Haji Yahya bin Begawan Mudim Dato Paduka Haji Bakar, the
Minister of Industry and Primary Resources, Brunei Darussalam for his interest and continuing
support of the botanical survey in Brunei. Special thanks are due to Hajah Nomiah Suria Hayati
binti Pehin Jamil (Permanent Secretary to the Ministry of Industry and Primary Resources),
Hajah Hasnah binti Haji Ibrahim (Deputy Permanent Secretary), Haji Saidin Salleh (Director
of Forestry), and Hajah Jamilah binti Haji Abdul Jalil (Acting Deputy Director of Forestry), as
well as Mahmud bin Haji Yussof, Chief Executive Officer of Brunei’s Heart of Borneo Centre.
We are likewise grateful for the mterest of past and present High Commissioners of Singapore
to Bmnei Damssalam, HE Joseph Koh and HE Jaya Ratnam, for facilitating the start of this
collaboration, which is jointly funded by both the Brunei Forestry Department and the National
Parks Board, Singapore. On the Singapore side, we thank the Ministry of National Development
and National Parks Board (NParks), in particular Kenneth Er, CEO of NParks; Dr Leong Chee
Chiew, Deputy CEO of NParks; Dr Nigel Taylor, Director of the Singapore Botanic Gardens;
and Dr David Middleton, Deputy Director and Keeper of the Singapore Herbarium (SING).
Many others have been most supportive and helpful to our programme: Roslinah binti Haji
Mohsin and Noralinda binti Haji Ibrahim (Forestiy Department, Brunei), Dr Kamariah Abu
Salim (Universiti Brunei Damssalam); Serena Lee, Paul K.F. Leong, Paul Pamsuraman Athen
and Alsa Moo (SING); Watu bin Awok, Jangamn anak Eri, Azlan bin Pandai, Muhanunad
Waffiuddin bin Zainol Ariffin, Mediana binti Kampong and Rauzaidi bin Mureh (BRUN). Not
least, the comments and suggestions of reviewers and the editor Dr David Middleton have
helped improve the text of this paper.

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MacKinnon, J. & MacKinnon, K. (1986). Review of the Protected Areas System in the Indo-
Malayan Realm. Gland: lUCN.

Merrill, E.D. (1921). A Bibliographic Enumeration of Bornean Plants. J. Straits Branch Roy.

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Merrill, E.D. (1950). A brief survey of the present status of Bomeon botany. Webbia 7: 309-
324.

Myers, N., Mittenneier, R.A., Mittermeier, C.G., da Fonseca, G.A.B. & Kent, J. (2000).

Biodiversity hotspots for conservation priorities. Nature 403(6772): 853-858.

Raes, N., Roos. M.C., Slik, J.W.F., van Loon, E.E. & ter Steege, H. (2009). Botanical richness
and endemicity patterns of Borneo derived from species distribution models. Ecography
32: 180-192.

Roos, M.C. (1993). State of affairs regarding Flora Malesiana: progress in revision work and
publication schedule. FI. Males. Bull. 11: 133-142.

Soepadmo, E. & Wong, K.M. (eds) (1995). Tree Flora of Sabah and Sarawak Volume 1.
Malaysia: Forest Research Institute Malaysia, Sabah Forestry Dept. & Sarawak Forestry
Dept.

Stibig, H.J. & Malingreau, J.P. (2003). Forest Cover of Insular Southeast Asia Mapped from
Recent Satellite Images of Coarse Spatial Resolution. Ambio 32 (7): 469^75.

Tan, H.S., Chung, R.C.K. & Soepadmo, E. (2011). A synopsis of Jarandersonia (Malvaceae -
Brownlowioideae). In: Wong, K.M., Leong-Skomickova, J., Lee, S. & Low, Y.W, (eds)
Proceedings of the 8th Flora Malesiana Symposium, Singapore, 23-27 August 2010.
Card. Bull. Singapore 63(1 & 2): 137-1 44.

Tate, R.B. (2002). The Geology^ of Borneo Island. CD-Rom. Kuala Lumpur: Geological Society
of Malaysia.

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Thiers, B. (continuously updated). Index Herbariorum: A global directory of public herbaria
and associated staff. New York Botanical Garden’s Virtual Herbarium. http://sweetgum.
nybg.org/ih/ (accessed on 31 May 2014).

Wilford, G.L. (1960). Geological Map of Brunei and Adjacent Parts of Sarawak. Scale
1:125,000. Sheets I and II. Kuala Lumpur: Geological Survey Department, Federation
of Malaya.

Wong, K.M. (1997). In Brunei Forests. An Introduction to the Plant Life of Brunei Darussalam,
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Darussalam: Forestry Department.

Wong, K.M. (1999). The nature of the Brunei flora. In: Wong, K.M. & Kamariah, A.S. (eds)
Trees and Forests of Brunei Darussalam. Pp. 53—73. Bandar Sri Begawan: University
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Bhd.

Wong, S.Y. & Boyce, P.C. (2011 ). Studies on Homalomeneae (Araceae) of Borneo X: Two new
Homalomena species from Brunei, and a new infonual species complex for supergroup
Cyrtocladon. Webbia 66{2)\ 133-141,

Wong, S.Y. & Boyce, P.C. (2014). Studies on Homalomeneae (Araceae) of Borneo XVIII:
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Gardens’ Bulletin Singapore 67(1): 61-68. 2015
doi: 10.3850/S2382581215000071

61

Novitates Bruneienses, 2. A remarkable new species of
Begonia sect. Petermannia (Begoniaceae)
from Brunei Darussalam

Y.W. Low\ A. A. Joffre^ & A.K. Muhammad Ariffin^

'Herbarium, Singapore Botanic Gardens, National Parks Board,

1 Cluny Road, 259569 Singapore
low_y ee_wen@nparks. go V. sg
^Bioinei National Herbarium, Forestry Department,
Ministry of Industry and Primary Resources, Jalan Menteri Besar,
Berakas, BB3910 Bmnei Damssalam

ABSTRACT. Begonia jamilahana Y.W.Low, Jofffe & Ariffin is described and illustrated
here as a new species based on a collection from Ladan Hills Forest Reserve, Tutong, Brunei
Damssalam. This new taxon is closely related to B. conniegeriae S. Julia & Kiew and B.
papyraptera Sands, but differs in a suite of morphological characters. Begonia jamilahana is
placed in Begonia section Petermannia, the largest of all the sections recognised.

Keywords. Begonia jamilahana, Begoniaceae, Borneo

Introduction

Begonia is a well-known genus, largely due to its popularity in horticulture. They
are mostly cultivated for their attractive foliage, as well as showy flowers (Holttum
& Enoch, 1995; Tebbit, 2005). At present, there are about 1500 species distributed
throughout the tropics, except in Australia, with many more taxa yet to be discovered
as individual species are commonly known to be confined to very small geographical
areas (Tebbit, 2005; Heywood et ah, 2007; Hughes, 2008).

Memll (1921), in his bibliographic enumeration, listed 26 species of Begonia
for Borneo, most of which were endemic species with the exception of B. repens
Blume. The checklist by Hughes (2008) enumerated 95 species for Borneo, all of
which are strictly restricted to the island. For Brunei, Coode et al. (1996) recorded 16
species, namely. Begonia awongii Sands, B. bahakensis Sands, B. baramensis Merr.,
B. bnmeiana Sands, B. chlorandra Sands, B. cyanescens Sands, B. eutricha Sands, B.
fuscisetosa Sands, B. hexaptera Sands, B. laccophora Sands, B. leucochlora Sands, B.
leucotricha Sands, B. papyraptera Sands, B. sibutensis Sands, B. stenogyna Sands and
B. tern burongen sis Sands. Four subspecies were recognised for Begonia bruneiana,
namely B. brimeiarm subsp. bruneiana, B. bruneiana subsp. angustifolia Sands, B.
bruneiana subsp. labiensis Sands and 5. bi^uneiana subsp. I'etakensis Sands. All of
the Brunei begonias were newly described by Sands (1996), with the exception of
Begonia baramensis Merr.

62

Gard. Bull. Singapore 67(1) 2015

A recent collection from the Ladan Hills Forest Reserve reveals yet another
novelty for Brunei from Begonia section Petermannia (Klotzsch) A.DC. as delimited
by Doorenbos et al. (1998). It has the following defining characters of the section,
namely (i) 2-tepaled male flowers, (ii) anthers with unilaterally positioned slits, (iii)
5-tepaled female flowers, (iv) inflorescence protogynous, with 1-2-flowered female
inflorescences, (v) trilocular ovai*y with axile placentation and bilamellate placentae,
and (vi) fruits with equal wings. The Ladan Hills novelty is closely related to Begonia
papyraptera, but differs in having (i) a smooth and terete stem in live material, (ii)
scantily scattered translucent hispid hairs on both sides of the leaf lamina, (iii) a mature
upper leaf surface that is dark green with red markings on vein axil nodes nearer to
the base of the leaf, (iv) a pale green lower leaf surface, and (v) base of capsule wings
cuneate. In contrast. Begonia papyraptera has (i) a ribbed and angular stem in live
material, (ii) a leaf lamina that is glabrous on both sides, (iii) a leaf upper surface that
is plain dark green with conspicuous white spots, (iv) a maroon lower leaf surface, and
(v) base of capsule wings rounded. Apart from Begonia papyraptera^ the Ladan Hills
taxon is also closely related to Begonia conniegeriae S. Julia & Kiew, but differs in the
characters enumerated in Table 1 . Hence, the Ladan Hills taxon is described here as
new.

New species

Begonia jamilahana Y.W.Low, Joffre & Ariffin, sp. iiov. (sect. Petermannia)

Similar to Begonia papyraptera Sands but differs in having a smooth and terete stem,
leaf blade scantily covered with franslucent hispid hairs on both sides, upper mature
leaf surface dark green with red markings on vein axil nodes nearer to the base of
the leaf, lower leaf surface pale green, and base of capsule wings cuneate. - TYPE:
Brunei, Tutong District, Lamunin, Ladan Hills Forest Reserve, Compartment 1, 58
m, 4°42’16”N 114°44’12”E, lowland mixed dipterocarp forest on yellow sandy clay
soils, steep slope, 15 Dec 2010, P. Azlan, A.K. Muhcl Ariffin, A. Watii & M. Rauzaidi
BRUN 23373 (holotype BRUN (including spirit material); isotypes K, SING). (Fig. 1,
2 )

Cane-like begonia, c. 40 cm tall, most parts of the plant glabrous except for the upper
and lower surface of lamina scantily covered with hispid translucent hairs neatly
arranged in between veins. Stems pale green, 0.8-1. 5 cm thick, very succulent, terete,
internodes 2.5-9 cm long, laxly branched, nodes red, not swollen. Stipules dark red,
narrowly lanceolate, c. 1-2 cm long, c. 0.3-0. 5 cm wide at the base, keeled, margin
entire, apex acute, caducous. Leaves alternate, oblique, distant, held horizontally;
petioles light green with both ends red, 5-20.5 cm long, terete; lamma dark green with
red markings on vein axil nodes nearer to the base of the leaf and iridescent above, pale
green beneath, young leaves very rarely with small white spots above, fading in older
leaves, asymmetric, elliptic to broadly elliptic, 12-19 x 11.5-14 cm, broad side 8-10.5

A new Begonia from Brunei Darussalam

63

cm wide, base cordate, basal lobes 3.5-7 cm, margin dentate, apex acuminate, acumen
1.5-2 cm long; venation palmate-pinnate, 7-9 veins, each branching dichotomously
2-3 times, vein nodes red near junction of petiole and gradually fading to pale green
towards the margin, flat and prominent on the upper surface, raised and prominent
on the lower surface. Inflorescences terminal, protogynous; female inflorescences
(l-)2 -flowered, positioned one node below the male inflorescences or solitary,
peduncles absent; male inflorescences composed up to 4 cymose partial inflorescences,
each monochasium with 3-6 flowers, peduncles 7-15 mm long. Bracts dark red, ovate,
keeled, c. 0.9-1. 1 cm long, c. 0.8 cm wide, margin entire, persistent. Male flowers:
pedicel white to pale pink, 7-9 mm long; tepals 2, pale pink with red flushes, ovate-
elliptic, 7-9 X 4-5 mm, margin entire, apex rounded; stamens c. 39, cluster conical,
stalked; filaments c. 0.8 nun long; anthers yellow, oblong, c. 1 mm long, apex slightly
emarginate. Female flowers: pedicel pale green, 4-6 mm long; ovary pale green, 1.3-2
X 0.3-0.4 cm, wings 3, equal, locules 3, placentae 2 per locule, placentation axile;
tepals 5, pink, outer 4 tepals ovate, c. 1 8-20 x 8-1 1 mm, innermost tepal oblanceolate,
c. 10 X 8 mm, margin entire, apex rounded; styles 3, deeply Y-shaped, c. 6 mm long,
divided c. 2 mm from the base; stigma yellow, papillose fonning a continuous twisted
band. Capsules (l-)2, pendent, 2.7-30 x 1,5-1 .7 cm, pale green when young, brown
when mature, locules 3, wings 3, equal, leathery when fresh, thinly fibrous when dry,
slightly expanded, tip rounded, dehiscing between locule and wing; pedicel c. 4-6 mm
long.

Additional specimens examined. BORNEO: Brunei: Tutong District, Lamunin, Ladan Hills
Forest Reserve, Ladan Hill, 20 Nov 2014, A.A. Joffre et al. BRUN 24029 (BRUN, SING).

Distribution and habitat. Begonia jamilahana is so far known only from the Ladan
Hills Forest Reserve, where it was documented on steep earth banks and adjacent
slopes in the mixed dipterocarp forest on yellow sandy clay soils.

Etymology. This species is named for Hajah Jamilah binti Haji Abdul Jalil, acting
Deputy Director of the Brunei Forestry Department, upon her retirement in 2015, as
a token of appreciation for her excellent service rendered to the Forestry Department,
as well as her unwavering support for the Brunei-Singapore botanical exploration
programme.

Provisional lUCN conservation assessment. Begonia jamilahana is so far known only
from a single population at Compartment 1, Ladan Hills Forest Reserve. The lUCN
status proposed here is Least Concern (LC) as the species is protected in a forest
reserve with no discernible threats. The Brunei Ministry of Industry and Primary
Resources officially announced in 2014 that logging will be phased out in all forest
reserves (Bmnei Times, 2014).

Notes. Begonia papyraptera Sands was described in Coode et al. (1996), based on a
single collection from the helipad Landing Point (LP) 286, Batu Apoi Forest Reserve,

64

Gard. Bull. Singapore 67(1) 2015

Table 1. Comparison of morphological characteristics, habitat and distribution among Begonia
conniegeriae S.Julia & Kiew, B. jamilahana Y.W.Low, Joffre & Ariffin and B. papyraptera
Sands.

Begonia conniegeriae

Begonia jamilahana

Begonia papyraptera

Stem (in life),
texture

Smooth and terete

Smooth and terete

Ribbed and angular

Stem (in life),
colour

Reddish or red brown

Pale green with red
nodes

Pale green

Stem nodes (in
life), swollen

Yes

No

Yes

Upper lamina

surface,

pubescence

Sparsely covered with
translucent erect hairs

Scantily covered with
translucent erect hairs

Glabrous

Upper lamina
surface (in life),
colour and
markings

Dark green

Dark green with red
markings on vein axil
nodes near tire leaf
base

Dark green with
white spots

Lower leaf

surface,

pubescence

Sparsely covered with
translucent erect hairs

Scantily covered with
translucent erect hairs

Glabrous

Lower leaf
surface (in life),
colour

Pale green

Pale green

Maroon

Capsule wings,
shape proportion

Unequal

Equal

Equal

Capsule wing
base, shape

Cuneate

Cuneate

Rounded

Habitat, substrate

Limestone

Yellow sandy clay
soils

Unknown

Distribution

Lobang Cina, Gunung
Mulu National Park,
Marudi District,
Sarawak, Malaysia

Ladan Hills Forest
Reseiwe, Tutong
district, Brunei
Damssalam

Batu Apoi Forest
Reserve, Temburong
district, Brunei
Darussalam

A new Begonia from Brunei Darussalam

65

Fig. 1. Begonia jamilahana Y.W.Low, Joffre & Arififin. A. Habit. B. Close-up of upper leaf
surface. C. Detail of stem showing red nodes. D. Close-up of male portion of inflorescence.
E. Close-up of female flowers. F. Detail of styles showing twisted papillose stigmas. G. Cross
section of an immature fruit. H. Immature fruit (side view). I. Detail of a stipule. A & B from
P. Azlan et al. BRUN 23373, C-J from ex-situ collection at BRUN. (Photos: A & B, A.K.
Muhammad Ariffin; C-J, Y. W. Low)

66

Gard. Bull. Singapore 67(1) 2015

Fig. 2. Distribution of Begonia conniegeriae S. Julia & Kiew, Begonia jamilahana Y.W.Low,
Joffre & Ariffin, and Begonia papyraptera Sands.

Temburong district {Johns et al. RJ7422), without a detailed description but with a
brief Latin diagnosis as follows:

“a B. congesta Ridley alls fructus latioribus, caulibus plusminusve glabris
(non ‘hirtis’), staminibus 15 vel plus (non 12), tepalis staminibus multo
longioribus (non brevioribus); a B. leucochlora (vide supra) fructus apice
rotundato (non alis acutis et fructus apice truncate), petiolis longioribus
differt.”

A total of three duplicates were gathered, with two sheets indicated as holotype in K
(Sheet 1 of 2: K000761105, and Sheet 2 of 2: K000761106); while the third sheet,
which is an isotype, is lodged in BRUN (herbaria acronyms follow Thiers, continuously
updated). For this study, the two holotype sheets were examined through the Southeast

A new Begonia from Brunei Darussalam

67

Asian Begonia database (Hughes & Pullan, 2007), as well as the JSTOR® Global Plants
website (Global Plants, continuously updated). Detailed examination was carried out
on the isotype preserved in BRUN. In addition, a living plant of Begonia papyraptera
was discovered recently by the second author in cultivation in the ex-situ collection of
begonias at the Kuala Belalong Field Study Centre (KBFSC). This cultivated specimen
in KBFSC matches the type materials represented in K and BRUN and, therefore,
provided us with further insights into the species for this study.

Begonia conniegeriae S. Julia & Kiew was described in Sang et al. (2013). We
have been unable to examine the materials as the isotype indicated in the article as having
been distributed to SING had not arrived by 13 March 201 5 (Serena Lee (SING), pers.
comm.). Images of the type materials were also not available for examination either in
the Southeast Asian Begonia database (Hughes & Pullan, 2007) or the JSTOR® Global
Plants website (Global Plants, continuously updated). The morphological characters of
Begonia conniegeriae used in this study are extracted from Sang et al. (2013).

ACKNOWLEDGEMENTS. We are grateful to Haji Saidin bin Salleh, the Director of Forestry
Department, Brunei Darussalam for providing us with field support and assistance for this study.
Special thanks to Hajah Jamilah binti Haji Abdul Jalil and Noralinda binti Haji Ibrahim for
their encouragement and continuous support and assistance. Dr Ruth Kiew (KEP) kindly made
available translated Latin diagnoses by Dr Mark Coode (K) of all the Begonia species described
by Martin Sands (K) published in Coode et al. (1996) for this study. Various staff members of
the Brunei National Herbarium (BRUN) have kindly helped: Mediana Kampong and Muhd.
Wafiuddin Zainol Arifhn assisted with herbarium materials and database; Watu Awok, Azlan
Pandai, Jangarun Eri and Rauzaidi Mureh provided excellent assistance in the field, as well
as maintaining the ex-situ reseai'ch collection. Research funding for YWL is provided by the
National Parks Board, Singapore and the Singapore Botanic Gardens. Christina Soh (Library of
Botany and Horticulture, Singapore Botanic Gardens) kindly helped with missing references.
We are grateful to the two reviewers, Dr Mark Hughes (E) and Dr Daniel C. Thomas (SING),
for their constructive comments in improving this manuscript. This research collaboration is
conducted under the MoU programme "'’The Botanical Sw’vey of Brunei Darussalam'’'' between
the Brunei Forestry Department, Ministry of Industry and Primary Resources and the National
Parks Board, Singapore.

References

Brunei Times (2014). Logging no longer allowed in forest reserves: Ministry of Industry
and Primary Resources. 23 March 2014. http://bt.com.bn/news-national/2014/03/23/
logging-no-longer-allowed-forest-reserves-mipr (accessed on 13 Feb. 2015).

Coode, M.J.E., Dransfield, J., Fonuan, L.L., Kirkup, D.W. & Said, l.M. (1996). A Checklist of
the Flowering Plants and Gymnosperms of Brunei Darussalam, xvii + All p. Brunei
Darussalam: Ministry of Industry and Primary Resources. (Begoniaceae, p. 38-41)

Doorenbos, J., Sosef, M.S.M. & de Wilde, J.J.F.E. (1998). The Sections of Begonia: including
descriptions, keys and species lists. 266 p. The Netherlands: Wageningen Agricultural
University.

68

Gard. Bull. Singapore 67(1) 2015

Global Plants (continuously updated). JSTOR® Global Plants. ITHAKA. http://plants.jstor.
org/ (accessed on 13 Mar, 2015).

Heywood, V.H., Brummitt, R.K., Culhani, A. & Seberg, O. (2007). Flowering Plant Families
of the World. 424 p. UK: Royal Botanic Gardens, Kew. {Begoniaceae, p. 58-59)

Holttum, R.E. & Enoch, I. (1995). Gardening in the Tropics, viii + 384 p. Singapore: Times
Edition.

Hughes, M. (2008). An Annotated Checklist of Southeast Asian Begonia, xi + 164 p. UK: Royal
Botanic Garden Edinburgh.

Hughes, M. & Pullan, M. (2007). Southeast Asian Begonia Database. Royal Botanic Garden
Edinburgh. http://elnier.rbge.org.uk/Begonia/ (accessed on 10 Dec. 2014).

Merrill, E.D. (1921). A bibliographic enumeration of Bornean plants. J. Straits Branch Roy.
Asiat. Soc. Special No. Singapore: Fraser & Neave Ltd. {Begoniaceae, p. 414-416)

Sands, M.J.S. (1996). Begoniaceae: Begonia. In: Coode, M.J.E., Dransfield, J., Fonnan, L.L.,
Kirkup, D. W. & Said, I.M. (eds) A Checklist of the Flowering Plants and Gymnosperms
of Brunei Damssalam. Brunei Darussalam: Ministry of Industry and Primary Resources.
{Begoniaceae, p. 38-41)

Sang, J., Kiew, R. & Geri, C. (2013). Revision of Begonia (Begoniaceae) from the Melinau
limestone in Gunung Mulu National Park and Gunung Buda National Park, Sarawak,
Borneo, including thirteen new species. Phytotaxa 99(1): 1-34.

Tebbit, M.C. (2005). Begonias: Cultivation, Identification and Natural History. Ill p. Portland:
Timber Press.

Thiers, B. (continuously updated). Index Herbariorum: A global directory of public herbaria
and associated staff. New York Botanical Garden’s Virtual Herbarium. http://sweetgum.
nybg.org/ih/ (accessed on 6 Mar. 201 5).

Gardens’ Bulletin Singapore 67(1): 69-84. 2015
doi: 10.3850/S2382581215000083

69

Novitates Bruneienses, 3. Eight new woody plants in the
Brunei flora, including flve new species

K.M. Wong & Y.W. Low

Herbarium, Singapore Botanic Gardens, National Parks Board,
1 Cluny Road, 259569 Singapore
wkin2000@gmai 1 .com

ABSTRACT. Five new species of seed plants are described: Alangium kayimiga K.M. Wong
(Cornaceae), Melastoma ariffinii K.M.Wong, Melastoma ashtonii K.M. Wong, Melastoma
jojfrei K.M.Wong & Y.W.Low (Melastomataceae), and Lasianthus jangarimii Y.W.Low
(Rubiaceae). New plant records for the ongoing Brunei floristic inventory include Planchonea
valida (Blunie) Blume (Lecythidaceae), Melastoma velutinosum Ridl. (Melastomataceae), and
Gardenia costiilata Ridl. (Rubiaceae).

Keywords. Borneo, Cornaceae, Lecythidaceae, Malesia, Melastomataceae, new records, new
species, Rubiaceae, Southeast Asia

Introduction

In this third installment of the series Novitates Bmneienses, we report our findings
of new species in four plant families. These include a new Alangium species
(Cornaceae), tliree new species and a new record in Melastoma (Melastomataceae), a
new Lasianthus species and a new Gardenia record (Rubiaceae), and a new record of
Planchonea (Lecythidaceae). Herbarium acronyms used follow Thiers (continuously
updated). All conservation assessments follow the methodology of lUCN (2012). For
the new species the assessments are global, for the new records we provide national
assessments for Brunei. All specimens cited have been seen by the authors.

Five new species

CORNACEAE

Alangium kayuniga K.M.Wong, sp. nov.

This new species is distinctive among the tree species of Alangium with a pinnately
veined leaf base, differing from A. havilandii Bloemb. in its subtruncate calyx tube
(the calyx tube in A. havilandii has distinct triangular teeth) and from A. javanicum
(Blume) Wangerin in its thicker leaves that dry medium brown, sparse and reticulate
tertiary venation and conspicuously asymmetric leaf base {A. javanicum has thinner
leaves typically di 7 ing olive brown to purple brown, regularly conspicuous scalarifonn
tertiary venation and typically symmetric leaf base). The new species also differs by its

70

Card. Bull. Singapore 67(1) 2015

glabrous calyx and fruit from both Alangium havilandii and A. javanicum, which have
a velvety hairy calyx and fruit. - TYPE: Brunei, Belait, Sungai Liang, Andulau Forest
Reserve, trail entrance to Compartment 7, mixed dipterocarp forest, 21 March 2007,
YusopBRUN 22161 (holotype BRUN; isotype SING). (Fig. 1)

Small tree 10-12 m tall, trunk 8 cm diameter. Leaves elliptic, 6.5-17 cm long, 2. 7-6. 6
cm wide, base cuneate, asymmetric, apex cuspidate (cusp 5-1 2 mm long), coriaceous,
glabrous, drying medium brown; midrib sunken on upper surface, prominent on lower
surface, secondary veins 7-9 pairs, tertiary venation sparse and reticulate; petioles 4-7
mm long, 1-1 .5 mm diameter, slightly channelled on upper side. Flowers not known.
Infructescence a short cyme, peduncle 3-6 mm long, with l-3(^) faiits. Fruits on
short pedicels 4-5 mm long, 1-1.5 mm diameter; ovoid-ellipsoid, slightly compressed
with 10 longitudinal ridges and grooves, 1.8-2. 2 cm long, 1.2-1. 4 cm wide; glabrous,
crowned by the persistent calyx limb and disc; calyx tube subtruncate, 1 .5-2 mm high,
glabrous. Seeds one per fruit, ellipsoid-compressed, 1.2-1 .3 cm long, 0. 8-0.9 cm
wide, 2-3 mm thick, testa smooth.

Etymology. ‘Kayu’ refers to tree or wood in Brunei Malay; this species is named for
Mohd. Niga bin Abdullah Nangkat, formerly Senior Forestry Assistant with the Brunei
National Herbarium and now retired, who led many collecting trips in Brunei forests.

Distribution. This would seem to be a very rare and narrow endemic in Brunei as it is
only known from the type collection.

Provisional lUCN conseivation assessment. Data Deficient (DD) as it has only been
collected once and no information is available on its distribution or population size.
Further field observations would enable a better understanding of its conservation
status in Bmnei.

Additional specimens examined. The species is only known from the type.

Notes. The slightly compressed ovoid-ellipsoid fruit, with longitudinal ridges and
grooves and crowned by the persistent calyx limb and disc, is characteristic oi Alangium
(Bloembergen, 1935, 1939). The most recent revision of Bornean Alangium was by
Berhaman (1995), in which ten species were recognised for the island. Of these, two
are climbers and another six have 3-5-veined leaf bases, characters not found in our
new species. Instead, it more closely resembles Alangium havilandii (with asymmetric
leaf bases and a swamp habitat) and A. javanicum (with symmetric leaf bases and a
mixed dipterocarp forest provenance). However, the glabrous calyx and fruit surface
oi Alangium kayuniga is consistent for both younger and more mature fruit, whereas in
A. havilandii and A. javanica the velvety nature of the calyx and fi'uit surface is evident
from young stages and persistent through to mature fruit.

Woody novelties in Brunei flora

71

Fig. 1. Alangium kayuniga K.M.Wong, fruiting branch. (Photo: Muhammad Ariffin)

MELASTOMATACEAE
Melastoma arijjinii K.M.Wong, sp.nov.

This new species resembles Melastoma stenophyllum Merr. in having linear leaves and
broad-triangular hypanthium scales that are less than three times as long as their width,
but differs in its hypanthium scales that are subulate and basally inflated with subentire
to sparsely denticulate margin, and which are more sparsely set and intermixed with
minute similar scales (in contrast to the irregularly short-fringed flat hypanthium scales
that are of one general size and closely overlapping in M. stenophyllum). - TYPE:
Tutong, Ramba, Ulu Tutong, down valley to SW of helicopter pad LP 239, 4°25'N
114°50'E, 150-200 m asl, 8 May 1992, Johns, Niga, Shanang & Han 7547 (holotype
BRUN; isotype K). (Fig. 2A-B)

Melastoma polyanthum var. linearifolium Bakh.f., Meded. Bot. Mus. Herb. Rijks
Univ. Utrecht 91: 69 (1943). - TYPE: Borneo, Gunong Narik, Kelam, May 1894,
Molengraaff B3460 (lectotype L, designated here).

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Fig. 2. Hypanthium scale types and detailed view of upper leaf surfaces in Melastoma ariffinii
(A, B), M ashtonii (C, D) and M joffrei (E, F). A & B from Othman et al. S 56078; C & D from
Ashton BRUN 5629; E & F from Wong & Joffre WKM 3207 (all SING). (Photomicrographs:
Y.W. Low)

Rheophytic bush 1-1.5 m high or more. Branch internodes covered with dense,
appressed, ovate-lanceolate, subentire to slightly short-serrate scales. Leaves with
petioles 0.4-2 cm long, c. 0.1 cm diameter; blades linear, 3.2-10 cm long, 0.4-1. 5 cm
wide, the 3 longitudinal veins sunken on the upper surface and prominent on the lower
surface, covered by a mixture of larger lanceolate appressed scales (c. 1 mm long)
and smaller appressed lanceolate scales (<0.3 mm long), lamina on the upper surface

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73

in dried material with abundant conspicuous elongate spicule-like patterns fomied by
rows of pale crystalliferous cells, glabrous or occasionally with micro-hairs less than
0.1 mm long (Fig. 2B), on the lower surface with scabrid scales with emergent scabrid
tips to 0.1 mm long. Flowers 1-3 in short cymes, peduncle 3-5 nun long, branches 1-2
mm long only; pedicels 4-6 mm long; hypanthium (Fig. 2 A) 5-7 imn long, 4-6 mm
diameter, green then magenta, with sparsely airanged scales, the scales subulate and
basally inflated, to 2 mm long, with entire to sparsely denticulate margins, intermixed
with minute similar scales mostly 0.5-1 nun long or shorter, lobes 5, broad-triangular,
5-6 mm long, alternating with inter-sepalar protuberances 1-2 nuu long; petals
obovate, c. 2.5 cm long, 1.8 cm broad, mauve; stamens not seen. Fruits c. 9 mm long,
8 mm diameter, green to reddish brown.

Etymology. The species is named after Muhammad Ariffin A. Kalat, experienced forest
botanist at the Brunei National Herbarium.

Distribution. This is apparently the common rheophytic Mela stoma species in
Clearwater streams of the Northwest Borneo region, including Brunei, Sarawak and
W Kalimantan.

Provisional lUCN conservation assessment. Least Concern (LC) as the species is
common and widespread. Besides that, the habitat in Brunei is also protected in Forest
Reserves and a National Park.

Additional specimens examined. BRUNEI: Temburong: First big waterfall on R. Temburong,
c. 500 ft, 6 Nov 1959, Ashton BRUN 759, (BRUN, K, SfNG). MALAYSIA: Sarawak: 5th
Division, Ulu Lawas, Kota Forest Reserve, on bank of Sg. Kota, 21 Oct 1971, Chai & Ilias S
31110 (A, K, L, SAN, SAR, SING); Kapit Division, Balleh, Ulu Mengiong, Sg. Entejum, 27
Oct 1988, Othman, Rantai & Jugah S 56078, (K, KEP, L, SAR, SING); Kapit Division, Sg.
Belaga at upper rapids, 12 Apr 1963, Ashton S 18242 (K, L, SAR, SING).

Notes. Meyer (200 1 ) had apparently not studied specimens of this rheophytic Melastoma
taxon in Borneo; none bears his annotation or are mentioned or indexed in his revision.
He placed Melastoma polyanthum var. linearifolium Bakh.f. in the synonymy of M.
malabathricum, but the variety has not hitherto been lectotypified. The syntypes are
MolengraaffB3460 from Mount Kelam in W Kalimantan (L) (which is our M. ariffinii
here) and Seimund s.n. from Kwala Teku in the Malay Peninsula (K: K000867833)
(which is a nan‘ow-leafed rheophytic foiua of M. malabathricum). Ridley’s Melastoma
polyanthum var. angustifolium (Ridley 1922: 765, as ‘angustifolia’) was applied
to this Malay Peninsula taxon and requires lectotypification (to be addressed in a
separate review of the Malayan species), but in any case is different from our new
species here. Here we lectotypify Melastoma polyanthum var. linearifolium with the
Bornean specimen, as Bakhuizen’s work addressed mainly the Netherlands East Indies
(Bakhuizen van den Brink Jr., 1943).

The Sumatran rheophyte Melastoma stenophyllum was also reduced by Meyer
(2001) to synonymy under M. malabathricum L. although it is, in our view, quite

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distinct: not only because the leaves are stenophyllous in an extreme manner, with
blade lengths 9-11 times the width (much more linear than narrow variations of the
typically elliptic leaves in M. malabathricum which has blade lengths 5-7(-8) times
the width), but also the flowers are typically solitary (compared to cymes of several to
7-9 flowers iiiM malabathricum).

Steenis (1981) confused ariffinii withM borneense (Cogn.) Bakh.f.,

which is typified by a collection from West Kalimantan: De Vriese 168 (L). The De
Vriese collection has leaves just L3-2.2 cm wide, but still clearly elliptic in shape,
and not sufficiently linear as in the truly rheophytic taxa; Melastoma ariffinii here
has leaves that are linear and only 0.4-1. 5 cm wide (length 8-10 times the width).
Also, the De Vriese specimen has 2-7 flowers in a cyme, and a hypanthium with
short-fringed, flat ovate-triangular scales, whereas Melastoma ariffinii has only 1-3
flowers and hypanthium scales that are subentire to remotely denticulate, and subulate
inflated structures. Lastly, the upper leaf surface has scabrid hairs 0.3-0. 5 mm long in
Melastoma borneense but subglabrous with only occasionally scabrous micro-hairs
less than 0. 1 mm long in M. ariffinii, so that the latter’s upper leaf surface is not scabrid
to the touch. We agree with Meyer (2001) that Melastoma borneense is synonymous
withM malabathricum', the two taxa have the same hypanthium scales and leaf upper
surface characters, among others.

Melastoma ashtonii K.M.Wong, sp.nov.

This new species resembles Melastoma stenophyllum Men*, in having linear leaves,
but differs in its hypanthium scales that are basally thickened and deeply incised into
linear segments (Fig. 2C) (in contrast to the fringed flat ovate-triangular scales on the
hypanthium in M. stenophyllum). - TYPE: Brunei, Belait, Sg. Ingei, rocky river bank,
21 January 1959, Ashton BRUN 5629 (holotype BRUN; isotypes K, L, SAR, SING).
(Fig. 2C-D)

Rlieophytic bush 1-2 m high, stems sometimes gnarled. Branch internodes covered
with dense, appressed, ovate-lanceolate, subentire to slightly short-serrate scales.
Leaves with petioles 0.5-1 .4 cm long, c. 0. 1 cm diameter; blades linear, 3.5-9 cm long,
0.4-1 cm wide, the 3 longitudinal veins sunken on the upper surface and prominent
on the lower surface, covered by a mixture of larger lanceolate scales 0.5- 1.2 mm
long and smaller lanceolate scales 0.2-0. 3 mm long, lamina on the upper surface in
dried material with abundant conspicuous pale elongate rows of crystalliferous cells
fomiing spicule-like patterns and quite glabrous (Fig. 2D), on the lower surface with
scabrid scales with 0.2-0. 3 mm emergent scabrid tips. Flowers 1-3 in short sessile
cymes, branches 1-2 mm long only; pedicels \-A mm long; hypanthium 10-11 mm
long, 7-8 mm diameter, green then crimson, moderately to densely scaly, the scales
with an entire thickened base 0. 1-0.2 mm high and linear segments 0.6-1.4(-2) mm
long, lobes 5, narrowly triangular, 8-10 mm long, alternating with inter-sepalar lobes
4-5 mm long; petals obovate, c. 3.3 cm long, c. 2 cm wide, mauve; short stamens
with a filament c. 7-8 mm long and anthers c. 5 mm long and 2.5-3 mm wide, with

Woody novelties in Brunei flora

75

apiculate tips, long anthers only one seen, filament unknown, anther c. 12 mm long.
Fruits (young) c. 8 mm long, 4 mm diameter, green to reddish brown.

Etymology. The species is named after Professor Peter Shaw Ashton, Brunei’s first
Forest Botanist who collected the type on his first assignment to the tropics, and who
later taught at Aberdeen and Haiward Universities.

Distribution. Known only from the Sungai Ingei area in Brunei.

Provisional lUCN conset'vation assessment. Least Concern (LC) as the habitat of the
species is well protected in the Sungai Ingei Conservation Area, which is also now
included within the designated Heart of Borneo area within Brunei.

Additional specimens examined. BRUNEI: Belait: Falls just upstream from Batu Melintang, 4
Jan 1989, Wong WKM 680 (BRUN, K, SING).

Notes. See comments under Melastoma ariffinii above. The new rheophytic species
here, Melastoma ashtonii, is also clearly different from M. malahathricum by its
hypanthium scale type: basally thickened scales that are deeply incised into linear
segments, often resembling miniature rakes, instead of the iiregularly short-serrate to
shallowly laciniate flat narrowly triangular-lanceolate scales as m M. malabathricum
(Fig. 3). Also, in both Melastoma malabathricum and M. stenophyllum, the pale
spicule-like patterns formed by rows of crystalliferous cells embedded in the upper leaf
surface are distally continuous with short scabrid hairs, but the upper laminar surface in
M. ashtonii is quite glabrous. In addition, Melastoma ashtonii has rather conspicuous
inter-sepalar lobes 4-5 mm long, compared to the much less conspicuous ones that
occasionally occur in M. malabathricum, absent altogether inM. stenophyllum.

Steenis (1981: 283) had already noticed the potential novelty of this taxon,
which he enumerated as “Melastoma sp. (nov. ?)” in his Rheophytes of the World.

Melastoma joffrei K.M.Wong & Y.W.Low, sp.nov.

This species is unique among Melastoma species with isomorphic stamens by
its hypanthium covered by small triangular scales of different sizes (Fig. 2E) (not
hairlike bristles as in M. cynoides Sm. and M. moluccanum Blume, nor penicillate
emergences as in M. montanum (Lauterb.) K.Meyer), very small bracts that do not
enclose the hypanthium (unlike in M. montanum), and scabrid upper leaf surfaces with
a mixture of longer and tiny coarse suberect to upcurved hairs (Fig. 2F) (not stidgose to
appressed pilose upper leaf surfaces as in the other species). - TYPE: Brunei, Tutong
district, Rambai, Ladan Hills Forest Reserve, on ridge NE of campsite beside Sungai
Buing, along old logging track, 13 October 2012, Wong & Joffre WKM 3207 (holotype
BRUN; isotype SING). (Fig. 2E-F, 4)

Treelet (single-stemmed) or shrub (with several stems from the base) to 1.5 m high.
Branch internodes eoarse with dense, spreading to erect small triangular scales.

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Gard. Bull. Singapore 67(1) 2015

Fig. 3. Consistently narrowly lanceolate, serrate-laciniate and densely overlapping hypanthium
scales, and scabrid-hairy upper leaf surfaces in Melastoma malabathricum from the Malay
Peninsula (A, B), Sabah (C, D) and Sarawak (E, F). A & B from Jagoe s.n.,C&.T) from Kadir
A 2855; E & F from Moulton 6723 (all SING). (Photomicrographs: Y.W. Eow)

Leaves with petioles 0.6-2 cm long, with a mixture of scattered to dense suberect short
scales and scabrid hairs; blades ovate-elliptic, 5.5-12.5 cm long, 1.5-4. 5 cm wide,
longitudinal nerves 3-5 (-7), sunken on upper surface, raised on lower surface and
bearing a mixture of suberect short scales and scabrid hairs, transverse veins between
the nerves scalariform, slightly sunken on upper surface, raised on lower surface;

Woody novelties in Brunei flora

77

Fig. 4. Melastoma joffrei K.M.Wong & Y.W.Low. Open flowers showing isomorphic stamens.
Inset: Detail of upper leaf surface showing different hair types contributing to a sandpapery
roughness. All from Wong & Joffre WKM 3207. (Photos: K.M. Wong)

upper laminar surface with a mixture of longer and tiny coarse suberect to upcurved
hairs, scabrid-sandpapery to the touch; lower laminar surface similarly scabrid.
Inflorescence a compact terminal cyme of (7-)9-13 flowers, main cyme branches
3-11 mm long, bracts tiny, broad triangular to semicircular, c. 1 mm long and wide.
Flowers small, only c. 2 cm across; hypanthium urceolate, 4.5-5 mm long, 4.5-5 mm
wide, surface scales short triangular, appressed, margins minutely denticulate, not or
only slightly imbricate and not entirely obscuring the hypanthium surface, green; calyx
lobes 5, broadly triangular, 1.5-2. 5 mm long, 1-2 mm wide; petals 5, obovate, 6-7.5
mm long (8-9 mm in fresh material), 4-5.5 mm wide (5-6 mm in fresh material),
the apex subtruncate to rounded, margins short-ciliate, pale purple-pink; stamens
10, isomorphic, filaments 3-4 mm long (3 .4-4.5 mm in fresh material), white, each
apically bearing two knoblike pale yellow appendages at the insertion of the anther,
anthers erect, 2-2.5 mm long (2.5-3 mm in fresh material), pale creamy yellow, apex
rounded, opening by apical pores; style slender-cylindric, 4. 5-5. 5 mm long (5.5-7
mm in fresh material), with a pale green base, then pink for the most part, apically
pale yellow; stigma rounded to subtruncate, 4-5-lobed, pale green. Fruit urceolate to
subglobose, 5.5-6 mm long, 5.5-6 mm wide, dehiscing irregularly transversely.

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Gard. Bull. Singapore 67(1) 2015

Etymology. The species is named for our colleague Joffre bin Haji Ali Ahmad, Forest
Botanist at the Brunei National Herbarium and experienced and long-time collector of
Brunei plants.

Additional specimens examined. BRUNEI: Tutong; Rambai, Ladan Hills Forest Reserve, Sg.
Buing, disturbed mixed dipterocarp forest, on ridge, 29 Oct 2013, Ariffin, Low, Y.W. & Azlan
BRUN 23229 (BRUN, SING). MALAYSIA. Sarawak: Tatau district, Anap, Muput Kanan,
Ulu Naoung, 6 m asl, 14 Oct 1963, Ashton S 19569 (A, BO, K, KEP, L, SAN, SAR, SING).

Distribution. Known only from the type locality in Brunei and another collection in
Sarawak’s Tatau district.

Provisional lUCN conservation assessment. Data Deficient (DD) as it has been only
twice collected with one of the collections made over 50 years ago. Field observations
are badly needed for a better understanding of its conservation status.

Notes. In Meyer’s revision of Melastoma (Meyer, 2001), there is a small group of
three Melastoma species with isomorphic stamens, M. cynoides, M. moluccanum and
M. montanum, which he recognises for the more seasonal parts of insular SE Asia and
New Guinea; the rest of the species in the genus have fiowers with dimorphic stamens.
Melastoma joffrei is Borneo’s first known tom a species with isomorphic stamens.

RUBIACEAE

Lasianthus jangamnii Y.W.Low, sp. nov.

This new species resembles Lasianthus linearifolius H.Zhu in having linear leaves but
differs by having triangular stipules, bullate leaves with a thin-papery and crispy texture,
a revolute margin and brochidodromous venation (with 17-40 pairs of secondary
veins fonning a distinct marginal vein), and a calyx with shorter tube (c, 0.4 mm long)
and shorter lobes (c. 0.4-0.9 mm long). In contrast, Lasianthus linearifolius has awl-
shaped (subulate) stipules, leaves with a plane surface and leaf margin, subcoriaceous
texture, craspedodromous venation (with 10-16 pairs of secondary veins), and a
calyx with longer tube (c. 1 mm long) and longer lobes (c. 1 mm long). - TYPE:
Brunei, Temburong, Amo, Sungai Temburong and Sungai Machang junction, ridge,
120-250 m alt., 18 September 1990, Puff et al. 900818-1/1 f mature fruits (holotype
BRUN; isotypes K, SAR, SING 2 sheets). (Fig. 5)

Understorey treelet c. 2 m high, stem c. 3^ mm wide. Branches solitary, slender,
terete, c. 1-2 mm wide, sparsely puberulent to subglabrous. Stipules narrowly
triangular, c. 1-3 mm long, sparsely puberulent. Petiole 4— 6(-7) mm long, 0.8-1. 2
mm wide, sparsely puberulent. lamina linear, (8.5-)l 1-13.7(-18.4) cm long,
(0.8-)l-1.5(-2.2) cm wide, bullate, thin-papery and crispy; leaf base cuneate; leaf
apex long-caudate; leaf margin entire to slightly repand, revolute; midrib prominent

Woody novelties in Brunei flora

79

Fig. 5. Lasianthus jangarunii Y.W.Low. A. Branches with conspicuously corrugated leaves.
B. Close-up of a flower bud and an immature fiaiit with persistent calyx lobes visible. C. Habit (Ulu
Temburong National Park). All from Low et al. LYW 629. (Photos: A-B: K.M. Wong, C: Y.W. Low)

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Gard. Bull. Singapore 67(1) 2015

and glabrous on both sides; secondary veins 17^0 pairs, making an angle of 85-90°
with the midrib and joining to fonn a distinct marginal vein, inconspicuous to slightly
raised and glabrous on upper side, prominent and glabrous on lower side; tertiary
venation inconspicuous to slightly raised on upper side, inconspicuous on lower side.
Inflorescences sessile; bracts absent. Flowers fascicular, sessile; calyx campanulate,
tube c. 0.4 mm long, outside surface puberulent; marginal lobes 4(-5), triangular,
c. 0.4-0.9 mm high; corolla hypocrateriform, white; tube c. 1 mm long, outside
puberulent; lobes 4(-5), triangular, c. 1.2 mm long, c. 1 mm wide, valvate, outside
puberulent. Drupes subglobose, c. 2-3 mm long, 2.5-3 mm wide, 4(-5)-ridged when
dry, puberulous, maturing black {Kirkup et al. 898); calyx persistent at fruit apex, with
4(-5) small triangular calyx lobes. Pyrenes 4(-5), thick-walled, each with a single
seed.

Etymology. This species is named for Jangarun anak Eri, knowledgeable field assistant
and tree climber attached to the Brunei National Herbarium (BRUN), who has assisted
numerous botanists with fieldwork.

Distribution and habitat. Lasianthus jangarunii is endemic to northwest Borneo
(Brunei: Temburong district, and Sarawak: Marudi district), where it grows on clay-
rich sedimentary soils as an understorey treelet on steep slopes of mixed dipterocarp
forest.

Provisional lUCN conservation assessment. Least Concern (LC) as the species is
protected in the Ulu Temburong National Park in Brunei and, although only collected
twice, in Sarawak it is recorded from Pulong Tau National Park where the habitat is
also protected.

Additional specimens examined. BRUNEI: Temburong: Amo, Kuala Belalong (‘Kuala
Temburong’ on label), hill, 21 Jul 1988, Wong WKM 253, leafy branch (BRUN, K, SING),
Batu Apoi Forest Reserve, on slope along Sungai Engkiang, 20 Nov 1991, Hansen 1592, young
fruits (BRUN); Amo, Ulu Belalong, LP 382, ridge top, 22 Jan 1994, Kirkup et al. 898, young
fruits (BRUN); Amo, Ulu Temburong National Park, hill behind Lubok Umar, ridge, 18 May
2014, Low et al. LYW 629, flower buds (BRUN, K, L, SAN, SING). MALAYSIA. Sarawak:
Marudi, Pulong Tau National Park (western part), Ulu Sungai Baong, on steep slope, 93 1 m
asl, 10 May 2007, Sang et al S 98058, young ifuits (K, KEP, L, SAN, SAR, SING); ibidem,
trail along the ridges to Bukit Tenidan, 1 1 May 2007, Muliati et al. S 97911, fruits (KEP, SAR).

Notes. Davis in Coode et al. (1996) identified this species as a taxonomically distinct
entity of Lasianthus from Temburong, which was listed only as Lasianthus “sp. 7”.
Zhu et al. (2012) evidently overlooked Coode et al. (1996) and this species while
preparing for the revision of Malesian Lasianthus; he stated that only “a selection
of the Malesian collections in the herbaria BKF, KEP, MO, and SING” had been
consulted. A recent collection from Ulu Temburong National Park {Low et al. LYW
629) matched Lasianthus “sp. 7” of Davis (Coode et al., 1 996), and is distinct from all
other Lasianthus taxa recognised by Zhu et al. (2012). This is so far the only known
Lasianthus species in Malesia with strongly bullate and linear leaves.

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81

Table 1. Comparison of morphological characteristics and distribution between Lasianthus
jangarimii Y.W.Low and L. linearifolius H.Zhu.

Lasianthus jangamnii Lasianthus Uneanfolius

Stipules triangular
Leaves bullate

Leaf texture thin-papery and erispy

Leaf margin revolute

Secondary veins 17-30 pairs

Loop-veined secondary venation
(broehidodromous)

Calyx tube c. 0.4 mm long

Calyx lobes c. 0.4-0. 9 mm long

Restricted to northwest Borneo (Brunei:
Temburong district, and Sarawak: Marudi
district) on clay-rich sedimentary soils

Stipules awl-shaped (subulate)

Leaves smooth

Leaf texture subcoriaceous

Leaf margin plane

Seeondary veins 10-16 pairs

Secondary venation emerging
from the midrib running towards
and terminating at the leaf margin
(eraspedodromou s)

Calyx tube c. 1 nun long

Calyx lobes e. 1 mm long

Restrieted to Mount Kinabalu (Malaysia:
Sabah) on ultramafic soil

Lasianthus jangarunii differs fromZ. linearifolius, which also has linear leaves,
by its stipules, leaf texture and venation, and calyx dimensions (see diagnosis above).
The character-states are compared in Table 1 .

Apart from that, Lasianthus jangarunii is restricted to northwest Borneo
(Brunei: Temburong district, and Sarawak: Marudi district) on clay-rich sedimentary
soils, whereas L. linearifolius is endemic to Mount Kinabalu (Sabah) growing on
ultramafic soil.

Lasianthus has some consistent characters that are easily recognised, when
adequate material permits, such as paired axillary inflorescences and pyreniferous
blue fruits (as in the type L. cyanocarpus Jack, but there are some species with white,
red or black fruits). In addition, Lasianthus spp. are more typically treelets in the
forest understorey with only solitary (not paired) primary branches developing along
the vertical stem. This branching feature was not noted in Zhu et al. (2012) but is
nevertheless important and represented in more recent collections of the genus; a good
example was illusfrated for Lasianthus pedicellatus H.Zhu in Zhu et al. (2012: 72, Fig.
30).

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Gard. Bull. Singapore 67(1) 2015

New plant records for Brunei

LECYTHIDACEAE
Planchonea valida (Blume) Blume

BRUNEI. Temburong: Amo, Kuala Belalong, Field Study Centre, 24 Apr 1998, Joffre BRUN
19016 {mum).

This is a new tree genus and species record for Brunei, collected after the publication
of Coode et al. (1996).

Provisional lUCN Conservation Assessment. Data Deficient (DD) for Brunei as the
species is only known from a single collection from Kuala Belalong. Further field
observations are needed for a better understanding of its conservation status in Brunei.

MELASTOMATACEAE
Melastoma velutinosum Ridl.

BRUNEI: Belait: Labi, Labi Forest Reserve, Compt 49, Ulu Sungai Rampayoh, 19 May 2009,
Yusop BRUN 22636 (BREfN); Melilas, Ulu Ingei hotsprings, 4°08’N, 114‘’43’E, 20 m asl, 7
Mar 1 996, Joffre BRUN 1 7295 (BRUN, SING). Tutong: Rambai, Ladan Hills Forest Reserve,
Bukit Bedawan, northwest of LP 263, 4‘’29'33"N 1 14''48'52"E, 250 m asl, 28 Mar 1997, Joffre
BRUN 18147 (BRUN, SING).

This species is known for the Malay Peninsula and Borneo (Sabah and Sarawak)
(Meyer, 2001). The several collections here record it for Brunei for the first time.

It is a treelet (single-stemmed) or slmib (with several stems from the base) to 3
m high. Distinctive features include the branch interaodes that are hirsute with dense,
spreading to curved long bristles; ovate-elliptic leaves with 3-5(-7) longitudinal nerves
and an upper lamina surface that is appressed-pilose (hairs to 0.5 mm long), slightly
velvety to the touch; a compact terminal cyme of 3-7 small flowers (only 2 cm across
in dried material); and a hypanthium densely covered by slender bristles 3-5 mm long.

Provisional lUCN conservation assessment. Least Concern (LC) for Brunei as the
species is common. This species occurs throughout northwest Borneo.

RUBIACEAE
Gardenia costulata Ridl.

BRUNEI: Belait: Andulau, without date, Ashton A2865 (BRUN). Temburong: Bukit Patoi,
without date, Ashton 3970 (BRUN).

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83

Gardenia costulata Ridl. was first described by Ridley (1934) based on a single
collection (Beccari 1986, K, BM) from Sarawak. While reviewing Sundaland
Gardenia, Low (2010) enumerated additional materials of G. costulata, including
a collection for Sabah (Madius SAN 50094) and Kalunantan {Hallier B1285), most
of which had been erroneously identified as G. pterocaJyx Valeton in many herbaria.
Gardenia costulata has truncate stipules with a revolute margin, and thin-coriaceous
leaves with an acuminate to long-cuspidate apex and a pubescent lower leaf surface.
These vegetative features adequately pemiit its distinction from G. pterocalyx, which
also has tmncate stipules but with a plane margin, and thicker leaves with a typically
rounded apex and a glabrous lower leaf surface. Although the two Brunei specimens at
the Brunei Herbarium were without flower or fruit, it was possible to match these two
specimens to G. costulata based on the distinctive stipule character, the shape of the
leaf apex, the leaf texture, and the presence of pubescence. Apart from moiphological
characters, the kerangas vegetation at Bukit Patoi in Brunei where Ashton 3970 was
collected also matches the habitat known for other specimens of G. costulata.

Gardenia costulata is restricted to Borneo, and so far has only been recorded
from kerangas forest. The habitat of Ashton A2865 from Andulau is not known,
although the main habitat type recorded for Andulau is Mixed Dipterocarp Forest
(Ashton, 1964).

Provisional lUCN conservation assessment. Data Deficient (DD) for Brunei as the
species is only known from two localities, naiuely Andulau (Belait District) and
Bukit Patoi (Temburong District). Field observations are badly needed for a better
understanding of its conservation status in Brunei.

ACKNOWLEDGEMENTS. The Ministry of Industry and Primary Resources and the Forestry
Department, Brunei Darussalam, and the Singapore Botanic Gardens, National Parks Board,
Singapore, provided the opportunity and fiuiding for this work. For special field coordination
and assistance, we thank Noralinda binti Haji Ibrahim, Roslinah bind Haji Mohsin (Forestiy
Department), Joffre Haji Ali Ahmad, Muhd. Ariffin A. Kalat, Watu bin Awok, Jangarun anak
Eri, Azlan bin Pandai, Muhammad Waffiuddin bin Zainol Ariffin and Rauzaidi bin Mureh
(BRUN); Serena Lee, Paul K.F. Leong and Paul Parusuraman Athen (SING). We are grateful
to the keepers and curators of the BO, BRUN, L, SAN, SAR and SfNG herbaria for facilitating
study of specimens in their care.

References

Ashton, P.S. (1964) Ecological Studies in the Mixed Dipterocarp Forests of Brunei State.

Oxford Forestry Memoirs 25. Oxford: Clarendon Press.

Bakhuizen van den Brink Jr., R.C. (1943). A contribution to the knowledge of the
Melastomataceae occurring in the Malay Archipelago especially in the Netherlands East
Indies. Meded. Bot. Mus. Herb. Rijks Univ. Utrecht 91: 1-391.

Berhaman, A. (1995). Alangiaceae. In: Soepadmo, E. & Wong, K.M. (eds) Tree Flora of Sabah
and Sarawak, vol. 1. Pp. 5-14. Malaysia: Forest Research Institute Malaysia, Sabah
Forestry Dept. & Sarawak Forestry Dept.

84

Gard. Bull. Singapore 67(1) 2015

Bloembergen, S. (1935). The genus Alangium in the Netherlands Indies. Blumea 1: 241-294.

Bloembergen, S. (1939). A revision of the genus Alangium. Bull Jard. Bot. Buitenzorg 3, 16:
139-235.

Coode, M.J.E., Dransfield, J., Fonnan, L.L., Khkup, D.W. & Idris, M.S. (1996). A Checklist
of the Flowering Plants and Gymnospenns of Brunei Darussalam. All p. Brunei
Darussalam: Ministry of Industry and Primary Resources.

lUCN (2012). lUCN Red List Categories and Criteria: Version 3.1. 2nd ed. Switzerland, Gland
and UK, Cambridge: lUCN.

Low, Y.W. (2010). Taxonomy and Biogeography of Sundaland Gardenia (Rubiaceae). M. Sc.
thesis. Kuala Lumpur: University of Malaya.

Meyer, K. (2001). Revision of the Southeast Asian genus Melastoma (Melastomataceae).
Blumea 46: 351-398.

Ridley, H.N. ( 1 922). The Flora of the Malay Peninsula, Vol. 1. London: L. Reeve and Company.

Ridley, H.N. (1934). New Malayan Rubiaceae. J. Bot. 72: 274—275.

Steenis, C.G.G.J. van (1981). Rheophytes of the World. 407 p. The Netherlands: Sijthoff &
Noordhoff

Thiers, B. (continuously updated). Index Herbariorum: A global directory of public herbaria
and associated staff New York Botanical Garden’s Virtual Herbarium. http://sweetgum.
nybg.org/ih/ (accessed on 31 May 2014).

Zhu, H., Roos, M.C. & Ridsdale, C.E. (2012). A taxonomic revision of the Malesian species of
Lasianthus (Rubiaceae). Blumea 57: 1-102.

Gardens’ Bulletin Singapore 67(1): 85-94. 2015
doi: 10.3850/S2382581215000101

85

Hoya undulata (Apocynaceae, Asclepiadoideae),
a new myrmecophytic species from Borneo,
and typification of H. darwinii

S. Rahayu\ U. Meve^ & M. Rodda^

'Bogor Botanic Gardens, Indonesian Institute of Sciences,

Bogor, Indonesia
srirahayukrb@yahoo.com

^Department of Plant Systematics, University of Bayreuth,
95440 Bayreuth, Germany

^Herbarium, Singapore Botanic Gardens, National Parks Board,
1 Cluny Road, 259569 Singapore
rodda.michele@gmai 1 .com

ABSTRACT. In this paper we document the discovery of a new Hoya species from Borneo,
H. undulata S.Rahayu & Rodda. The leaves of the new species form domatia, specialised
structures harbouring ants, previously only observed in Hoya mitrata Kerr and H. darwinii
Loher. The corona is unique among Bornean Hoya species as its lobes develop two lateral
hooked appendages, also present in H. griffithii Hook.f, a species from mainland Asia. A
lectotype for Hoya darwinii is also selected.

Keywords. Ant plant, domatia, heath forest, Hoya mitrata, Indonesia, myrmecophyte. West
Kalimantan

Introduction

Hoya R.Br. is a poorly known tropical genus for which a modern revision is lacking.
Extensive field investigations and accurate documentation of new species are essential
before a generic revision can be compiled (Rodda & Ercole, 2014). In Borneo,
Merrill (1921) listed only eight Hoya species and more recently Forster et al. (1998)
described the diversity of the genus in Borneo as ‘unknown’. Recently, new taxa have
been described based on collections from Sabah (e.g. Lamb et al., 2014; Rodda et
al., 2013) and to a lesser extent, Sarawak (Rodda & Simonsson, 2011a, b; Rodda &
Simonsson Juhonewe, 2013a). The higher number of novelties described from Sabah
is directly linked to efforts by the staff at Kipandi Park (Kampung Kipandi, Moyog,
Sabah) to cultivate wild-collected sterile specimens until they bloom so that they can
be identified (Lamb et al., 2014), an approach successfully adopted by Rintz (1978) in
Peninsular Malaysia. Among the species described in Lamb et al. (2014) only Hoya
sammannaniana A.Lamb et al. is also found in Kalimantan. Two further species, Hoya
rintzii Rodda & Simonsson and Hoya beccarii Rodda & Simonsson can be found
throughout Borneo and are widespread elsewhere in SE Asia (Rodda & Simonsson

86

Gard. Bull. Singapore 67(1) 2015

Jiionewe, 2013b; Rodda et al., 2014). The diversity of Hoya in Indonesian Borneo is
least known due to the scarcity of recent collections; the only information on Hoya
diversity in Central Kalimantan can be found in Rahayu (2006) who lists only nine
species. Lamb et al. (20 14) suggested that 60 to 70 Hoya species may occur in Sabah
alone and it is becoming apparent that the diversity of Hoya of Borneo is expected to
be comparable to or exceeding that of the Philippines with 104 species (Aurigue et al.,
2013) or of New Guinea with 74 species (Forster, 1996).

During a recent expedition in West Kalimantan Mr Sulaiman Hasim discovered
a new Hoya species that we formally describe and illustrate here. It is compared with
the vegetatively similar Hoya darwini and H mitrata. The latter was lectotypified in
Rodda (2012) while the former is here lectotypified.

Hoya undulata S. Rahayu & Rodda sp. nov.

Similar to Hoya mitrata Kerr and H. darwinii Loher when sterile because it has
specialised convex leaves foiming multileaved domatia. Easily separated when fertile
because the flowers have a rotate corolla formed by almost completely free corolla
lobes, while Hoya mitrata and H. danvinii have reflexed corollas with a tube almost as
long as the lobes. - TYPE: Indonesia, West Kalimantan, Putussibau, 300-380 m, on a
slope above a stream, February 2014, Sulaiman Hasim s.n. (holotype BO). (Fig. 1, 2)

Epiphytic climber with white latex in all vegetative parts, glabrous. Stems slender,
intemodes very variable in length, from 0.5 cm in the parts ofthe vine forming domatia, to
20 cm long in climbing stems, 3-5 mm diameter, green with darker spots; adventitious
root sparsely produced along the stem. Leaves petiolate; petiole broader than stem,
0.5-2. 5 cm long, 3-5 mm in diameter, light green to purple; lamina dimorphic, the first
type ovate, obovate or oblanceolate, usually almost flat, located along climbing stems,
10-20 X 5-10 cm, the second type almost round to elliptic, usually convex and forming
domatia with 4-10 leaves each 3-7 (-10) x (2-)4-5 cm, base cordate with numerous
minute basal colleters fonning a convex line on the adaxial surface at the base of the
lamina, apex acute or acuminate, margins minutely undulate but appearing minutely
dentate in the proximal half of the lamina, sometimes entire along the distal part of the
lamina; venation pinnate, secondary veins 4-5 pairs, diverging at c. 70®, anastomosing,
tertiary venation reticulate; dark green above with grey to purple spots, below lighter
green, occasionally all purple, midrib depressed above, raised below, light green,
sometimes purple. Inflorescence positively geotropic, pseudo-umbellate, convex,
2-5-flowered(-12 flowered in cultivated material); peduncle positively geotropic
or ageotropic, l-2(-5) cm long, 3-5 mm in diameter, green with dark purple spots,
glabrous. Flowers unscented, lasting 4-5 days in cultivation; pedicel 2. 5^.5 cm long,
1.5-2 mm in diameter, white-light green with red spots, glabrous. Calyx lobes ovate,
2-2.5 X 1.5-2 mm, light pink to purple, glabrous with ciliate translucent margins,
basal colleters one in each calyx lobe sinus, 0.3-0. 5 x c. 0.3 mm, ovate to triangular
with a round tip. Corolla rotate, concave, with corolla lobe tips inflexed, 2,5-4 cm
in diameter, 4-5 cm when flattened; corolla lobes almost free, ovate to oblanceolate.

New Hoya from Borneo

87

Fig. 1. Hoya undulata S.Raha)^ & Rodda (from M. Rodda MR650, SING) A. Buds. B.
Inflorescence, top view. C. Inflorescence from underneath. D. Domatia; E. Leaf, abaxial, with
a magnification of the minutely undulate margin. F. Leaf, adaxial, with a magnification of the
basal colleters. (Photos: A, B, E, F, Michele Rodda; D, Surisa Somadee)

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Gard. Bull. Singapore 67(1) 2015

Fig. 2. Hoya undulata S.Rahayu & Rodda (from M. Rodda MR650, SING). A, B, C. Flower.
D. Corolla after removing the corona. E. Corona, top view. F. Corona, from underneath. G.
Corona, side view. H. Calyx and ovaries. I. Pollinarium with twin pollinia, with a magnification
of the much reduced sterile edge. (Photos: Michele Rodda)

New Hoya from Borneo

89

2-2.5 X 1-1.5 cm, white with purple red dots outside, cream-light pink inside, margin
recurved predominantly in the proximal half of the lobe, apex acute, curved toward the
adaxial surface, pilose inside with glabrous apex, glabrous outside. Corona staminal,
12-13.5 mm in diameter, 5-7 mm high; lobes spreading, when observed from above
clavate, 5. 5-6. 5 x 2.5-3 mm, carinate above, below sulcate with revolute margins,
outer process concave, inner apex rounded, upcurved, abruptly narrowing into a terete
linear inner process with upcurved acuminate tip; two lateral hooked appendages at
the junction between outer and inner process. Anthers ovate to almost round, c. 1.5 x
1.5 mm, with apical round membranaceous appendage just exceeding the style-head
apex. Pollinia oblong, 900-1000 x 350^00 pm, narrowing towards the base, apex
round, sterile edge much reduced, limited to the upper outer part of the pollinium;
corpusciilum rhomboid with acute tips, 500-600 x 200-250 pm; caudicle broadly
triangular, hyaline, unwinged, 250-300 x 250-300 pm at the widest. Style-head 5
angled in cross section, with 5 lobes alternating with the stamens, style-head apex
round, 1 .5 mm long, c. 3 mm broad at the base. Ovaries 2, conical with round tip,
1.5-2 mm long, c. 1 mm wide at the base, light green, red at the base and at the apex.
Fruit and seed not seen.

Distribution. Only known from the type locality in West Kalimantan, close to
Putussibau. A second unlocalised collection is widely available in cultivation.

Etymolog)?. The specific epithet refers to the characteristic undulate leaf margin.

Habitat and ecology. Observed in lowland heath forest at 300-380 m above sea level
on a slope above a stream, epiphytic on small tree trunks about 1.5 m above ground,
growing in about 80% humidity and 50% sunlight. (Sulaiman Hasim pers. comm.).
According to the epiphytic zonation by .Johansson (1975), the species grows in zone
B. The zonation of epiphytes is mainly correlated with their light, nutiients and water
requirement. Some species are restricted to strongly illuminated sites, some to shady
sites, while some avoid both strong light and deep shade and yet others have a wide
range of tolerance (Benzing, 1990). Usually, occupying the B zone means that the
species is adapted to moist shady habitats. The leaves of Hoya undulata can adapt to
high light levels, but they have been observed to develop a purple colour in intense
sunlight. At the type locality, the plants were rooting in ant nests and, on casual
observation, the leaf domatia harboured ants.

Provisional lUCN conservation assessment. Known from only one locality, the
preliminaiy conservation status of Hoya undulata is Data Deficient (DD, lUCN 2014).
Ex situ collections are present in Bogor Botanic Gardens (from the type locality) and
in Singapore Botanic Gardens (SBG acc. no. 20132428).

Notes. Two species, Hoya mitrata, from Thailand, Peninsular Malaysia, Sumatra,
Borneo, Sulawesi and Java, and H. darwinii, endemic to the Philippines, both
commonly associated with ants (Kleijn & Donkelaar, 2001), are similar to H. undulata.

90

Gard. Bull. Singapore 67(1) 2015

Fig. 3. Hoya mitrata Kerr (Photographed at Gunung Panti, Johor, Malaysia) A. Inflorescence.
B. A small domatium. Hoya darwinii Loher (M Rodda MR421, SING). C. Inflorescence. D.
Domatia. E. Leaves not forming domatia. (Photos: Michele Rodda)

In vegetative morphology, the three speeies have two types of leaves, the first with a
fiat or slightly eonvex lamina, occurring at widely spaced nodes on long climbing
stems, the other convex, occurring on stems with shorter intemodes, and these forming
domatia harbouring ant colonies. The first type of leaf is oblong-lanceolate in Hoya

New Hoya from Borneo

91

mitrata and broadly lanceolate in H. danvinii, both with an entire edge, while in H.
undulata it can be ovate, obovate or oblanceolate, with a minutely undulate margin
which may appear minutely dentate in the proximal half of the lamina, sometimes
entire along the distal part of the lamina. In Hoya danvinii the domatia are globose,
4-7 cm in diameter and fonned by 2-6 convex leaves with an inrolled edge (Fig.
3D). The domatia of Hoya mitrata are instead cabbage-shaped, usually larger, fonned
by tightly clustered broadly obovate convex leaves without inrolled edges (Fig. 3B)
(Weissflog et al., 1999; Kleijn & Donkelaar, 2001). In its natural environment, the
domatia of Hoya undulata are generally looser than those of//, mitrata or//, dai^vinii,
formed by 4-10, round to elliptic convex leaves 3-7 (-1 0) cm long, while in cultivation
domatia rarely develop and are usually formed by larger leaves (Fig. ID),

The inflorescences of Hoya darwinii and H. mitrata are flat to slightly convex,
negatively geotropic, bearing 10-15 flowers (Fig. 3 A, C), the corolla has a tube almost
as long as the lobes, and the lobes are reflexed; the outer process of the corona lobes
is swollen, erect. The inflorescences of Hoya undulata are also flat to slightly convex
but instead positively geotropic (Fig, lA-C), bear 2-5 flowers (exceptionally 12 in
cultivation); the corolla is concave, with almost free lobes, the corona lobes outer
process is concave, spreading with an upcurved apex.

The pollinia of Hoya darwnnii and H mitrata do not have a sterile edge, a
character that led Kloppenburg (1994) to accommodate both species in the new section
Rudimentalia Kloppenb. The pollinaria of Hoya undulata are overall similar to those
of //. mitrata and //. darwinii but its pollinia instead have a sterile edge, albeit much
reduced, limited to the upper outer side of the pollinium (Fig. 21).

A striking similarity can be observed between the corona of Hoya undulata and
H. griffithii from mainland Asia. The corona lobes are clavate in both species, with two
lateral hooked appendages at the junction between the outer and the inner processes.
These appendages have not been documented in any other Hoya species.

A key to separate Hoya danvinii, H. mitrata and H. undulata is provided below.

Other specimens examined. Unlocalised, 30 March 2014, Rodda, M. MR650, collected from
material cultivated at the Singapore Botanic Gardens, acc. no. 20132428 (SING).

Key to Hoya species forming domatia

la. Inflorescence negatively geotropic with 10-15 flowers, outer process of corona

lobes swollen, erect 2

lb. Inflorescence positively geotropic, with 2-5 (-12) flowers, outer process of corona

lobes concave, spreading with upcurved apex Hoya undulata

2a. Domatia globose, formed by concave leaves with inrolled edges
Hoya darwinii

2b. Domatia cabbage-shaped, formed by vaulted leaves without inrolled edges

Hoya mitrata

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Gard. Bull. Singapore 67(1) 2015

Lectotypification of Hoya darwinii

Hoya darwinii Loher, Gard. Chron. 47: 66 (1910). - TYPE: Philippines, Luzon, Rizal
Prov., September 1909, Loher, A. 14574 (lectotype UC [UC243291], designated here).

When Loher described darwinii he did not cite any specimens as belonging to the
taxon. He only indicated that the species was collected in Luzon and that it bloomed
in March and April. Loher collected in Luzon over a long period of time, from 1889
to 1915. His specimens collected before 1906 were deposited at K, with substantial
sets of duplicates in PNH, US, M, CAL; his collections from 1908 to 1915 (numbers
12000 to 15170) were deposited in PNH (Merrill, 1925), with a first duplicate set sent
to M and additional specimens to UC and A (orchids only) (Steenis-Kruseman, 1950).
We have examined Hoya specimens at K, and made enquires to M, UC, PNH and US
about the presence of any relevant Loher material. At K we located numerous early
Loher specimens, none identifiable as Hoya daminir, at M we found post- 1908 Loher
specimens, once again not belonging to H, danvinii. No Hoya darwinii specimens
collected by Loher could be found at PNH or US. However, three specimens of Hoya
danvinii were found at UC: two duplicates of Loher 14990, collected in Luzon, Rizal
Prov., in 1915, one of which is also labelled 395 (this label should be removed

as it likely refers to a different Hoya species with yellow flowers, while H darwinii
generally has pink flowers), and Loher 14574, collected in Luzon in September 1909
(Fig. 4). The latter is the only available specimen collected before the publication date
of Hoya danvinii and most probably belongs to the original material on which this
taxon is based. Thus, it is the only specimen available for lectotypification. However,
it must be noted that it does not fully match Loher ’s description as the flowering
period was indicated as March-April. If new evidence were to be found to indicate
this specimen is not original material, and in the absence of the discovery of alternative
original material, then the lectotypification designated here would count as an effective
neotypification under Art. 9.9 of the ICN (McNeill et al., 2012).

ACKNOWLEDGEMENTS. We would like to thank Mr Sulaiman Hasim who provided material
of the new species for study and cultivation in Bogor Botanic Gardens; Yoyo Budiman for his
assistance in transferring the material, Teguh (BBG) for assistance in pressing the materials and
Surisa Somadee, who provided photographs oi Hoya undidata. We also thank the curators of
BCU, BK, BKF, BM, BO, BRUN, FI, K, KEP, L, LAE, M, P, PNH, SAN, SAR, SNP, SING,
UC, US and UPM herbaria for allowing access and/or for providing high quality images of
herbarium specimens, an anonymous reviewer and Dr David Middleton, Editor of Gardens’
Bulletin Singapore, for suggesting improvements to this manuscript.

New Hoya from Borneo

93

References

Aurigue, F.B., Sahagun, J.R. & Suarez, W.M. (2013). Hoya cutis-porcelana (Apocynaceae): A
New Species from Samar and Biliran Islands, Philippines. J. Nat. Stud. 12 (1): 12-17.

Benzing, D.H. ( 1 990). Vascular epiphytes: general biology and related biota. Cambridge Univ.
Press, Cambridge..

Forster, P.l. ( 1 996). A checklist of the Asclepiadaceae of Papua. Sci. New’ Guinea 22(1 ): 1 5-22.

Forster, PL, Liddle, D.J., & Liddle, I.M. (1998). Diversity in the genus Hoya (Asclepiadaceae-
Marsdenieae). Aloe 35: 44^8,

Johansson, D.R. (1975). Ecology of epiphytic orchids in West African rain forests. Amer.
Orchid Soc. Bull. 44: 125-136.

Kleijn, D. & van Donkelaar, R. (2001). Notes on the taxonomy and ecology of the genus Hoya
{Asclepiadaceae) in Central Sulawesi. Bluniea 46: 457-483.

Kloppenburg, R.D. (1994). Hoya sections. Fraterna 1994(1&2): I-XXIV.

lUCN Standards and Petitions Subcommittee (2014). Guidelines for Using the lUCN Red
List Categories and Criteria. Version 10.1. http://www.iucnredlist.org/documents/
RedListGuidelines.pdf (accessed on 20 Jan. 2014).

Lamb, A., Gavrus, A., Emoi, B. & Gokusing, L. (2014). The hoyas of Sabah, a commentary
with seven new species and a new subspecies. Sandakania 1 9: 1-89.

McNeill, J., Buck, W.R., Demoulin, V., Greuter, W., Hawkworth, D.L., Herendeen, P.S., Knapp,
S., Marhold, K., Prado, J., Pmd’homme van Reine, W.F., Smith, G.F., Wiersema, .T.H. &
Turland, N.J. (2012). International Code of Nomenclature for algae, fungi, and plants
(Melbourne Code). Regnum Vegetabile 154. 205 p. Konigstein: Koeltz Scientific Books.

Merrill, E.D. (1921). A bibliographic enumeration of Bornean plants. J. Straits Branch Roy.
Asiat. Soc., Special Number. Singapore: Fraser & Neave Ltd.

Merrill, E.D. (1925). New species of Philippine plants collected by A. Loher. Philipp. J. Sci.
27: 21-59.

Rahayu, S. (2006). Keanekai'agaman jenis Hoya di Hutan Lindung Bukit Batikap, Pegunungan
Muler, Kalimantan Tengah. Biodiversitas 7(2): 139-142.

Rintz, R.E. (1978). The Peninsular Malaysian species of Hoya {Asclepiadaceae). Malayan Nat.
J. 30: 467-522.

Rodda, M. (2012). (2074) Proposal to conserve the name Hoya mitrata against H. wallichiana
{Apocynaceae, Asclepiadoideae). Taxon 61(3): 685-686.

Rodda, M. & Ercole, E. (2014). Hoya papaschonii (Apocynaceae: Asclepiadoideae), a new
species from southern Thailand with a peculiar corona. Phytotaxa 175(2): 097-106.

Rodda, M. & Simonsson, N. (2011a). Hoya devogelii {Apocynaceae: Asclepiadoideae), a new
species from Sarawak, Borneo. Webbia 66(1): 33-38.

Rodda, M, & Simonsson, N. (2011b). Hoya medinillifolia (Apocynaceae: Asclepiadoideae), a
new species from lowland forest of Sarawak, Borneo. Webbia 66(2): 149-154.

Rodda, M. & Simonsson Juhonewe,N. (20 13a). Hoya nuttiana (Apocynaceae: Asclepiadoideae),
a new species from Sarawak, Malaysian Borneo. Phytotaxa 140 (1): 56-60.

Rodda, M. & Simonsson Juhonewe, N. (2013b). The taxonomy of Hoya micrantha and Hoya
revoluta (Apocynaceae, Asclepiadoideae). Webbia 68(1): 7-16.

Rodda, M., Simonsson Juhonewe, N. & Ercole, E. (2013). Hoya coiymbosa (Apocynaceae:
Asclepiadoideae), a new unusual species from Sabah, Borneo and its systematic position
based on phylogenetic analyses. Syst. Bot. 38 (4): 1125-1131.

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Gard. Bull. Singapore 67(1) 2015

Rodda, M., Simonsson Juhonewe, N. & Rahayu, S. (2014). Taxonomic revision of the Hoya
mindorensis complex (Apocynaceae:-Asclepiadoideae). Webbia 69(1): 39^7.
Steenis-Kruseman, M.J. van (1950). Malaysian plant collectors and collections: Being a
cyclopaedia of botanical exploration in Malaysia and a guide to the concerned literature
up to the year 1950. Flora Malesiana, Ser. 1,1: 5-605. Jakarta: R Noordhoff
Weissflog, A., Moog, J., Federle, W., Wernr, M., Hashim, R. & Maschwitz, U. (1999). Hoya
mitrata Kerr. (Asclepiadaceae): a new myrmecotropic epiphyte from Southeast Asia
wih a unique multileaved domatium. Ecotropica 5: 221-225.

Gardens’ Bulletin Singapore 67(1): 95-99. 2015
doi: 10.3850/S2382581215000101

95

Begonia bimaensis, a new species of Begonia from
Sumbawa Island, Indonesia

N.K.E. Undaharta^, I Made Ardaka, Agung Kumiawan & Bayu Adjie

Bali Botanic Garden - Indonesian Institute of Sciences
Candilaining, Baturiti, Tabanan 82191, Indonesia
'nika002@lipi.go.id

ABSTRACT. A new species of Begonia sect. Reichenheimea, B. bimaensis Undaharta &
Ardaka, is described from Mt Muria in Binia District, Sumbawa Island, Indonesia. The species
is a narrow endemic, currently known from only one population, and has been assessed as
Vulnerable using lUCN criteria.

Keywords. Begonia, Sumbawa, new species

Introduction

The pantropical genus Begonia L. is one of the largest genera of flowering plants,
comprising more than 1700 species (Hughes, 2008 and the many species described
since then). Southeast Asia is a hotspot of species diversity for Begonia (Hughes,
2008), with Indonesia being particularly rich in species and with many species still
being discovered (Hughes, 2008; Thomas & Hughes, 2008; Girmansyah, 2009, 2012;
Girmansyah et ah, 2009; Hughes et ah, 2009; Thomas et ah, 2009, 2011; Ardi &
Hughes, 2010; Wiriadinata, 2011; Ardi et ah, 2013, 2014; Ardi, 2014; Lin & Peng,
2014). From these works the current estimate for the total number of Begonia species
in Indonesia is 213 species.

There are only a few Begonia species recorded for Nusa Tenggara Barat (West
Nusa Tenggara), and currently only two species have been reliably recorded from
the largest island, Sumbawa {B. multangula Blume - Hughes & Pullan, 2007; and B.
muricata Blume - Hughes, 2008). Examination of herbarium specimens shows that
several endemic species remain to be described from the region. The lack of records
for widely distributed species, such as Begonia longifolia Blume, may also reflect the
poor documentation of the Begonia flora of Sumbawa, since we have seen herbarium
material of this species from the sun'ounding islands of Bali, Lombok, Sumba and
Timor.

During a fieldtrip to Sumbawa Island, a small population of an unknown Begonia
from Mount Maria was found. Two living specimens were brought to the Bali Botanic
Garden where they were cultivated. As these specimens could not be identified, the
new species Begonia bimaensis is described here. Begonia bimaensis is placed in
Begonia section Reichenheimia (Klotzsch) A.DC. as it exhibits the characters typical
of the section: rhizomatous or tuberous stems, protandrous inflorescences, and three
locular ovaries with entire placentae (Doorenbos et ah, 1998).

96

Card. Bull. Singapore 67(1) 2015

All available Begonia specimens from the Herbarium of Bali Botanic Garden,
BO, E, K, L and SfNG have been consulted and hence it must be assumed, at least until
more intensive collecting in Sumbawa reveals otherwise, that the species described
here has a very restricted range.

Species description

Begonia bimaensis Undaharta & Ardaka sp. nov. § Reichenheimia
Similar to Begonia sendangensis Ardi but differs in the dense white, long hairs on
the adaxial surface of leaves (versus glabrous), female flower with four tepals (versus
three) and the ovary with 3 wings, reddish, equal or subequal, locules 3, placentation
axillary. Ovary characters for Begonia sendangensis, pinkish, equal, rounded at base,
placentae axile. - TYPE: Cultivated at Bali Botanical Garden from vegetative material
collected in the wild from Indonesia, West Nusa Tenggara, Sumbawa Island, Bima,
Wawo, Ntori, Mt. Maria, 08°29'55"S 118°52'41"E, 525 m elev., 4 October 2010.
Cultivated material vouchered and selected as type material on 11 December 2012
under I Made Suja HK 1276 (holotype Herbarium Bali Botanic Garden; isotype BO).
(Fig 1)

Perennial, tuberous, monoecious small herb, less than 10 cm tall. Stems very reduced,
an elongated tuber 1.5-1 . 8 cm long, diameter 3-4 mm, intemodes c. 1 mm long;
stipules ovate to elliptic, c. 3 x 1-2 mm, pale green, apex projecting up to c. 6 mm long,
persistent. Leaves alternate; petioles 3. 5-7. 5 cm long, red, sparsely covered with long
pilose hairs c. 2-5 mm long; lamina basifixed, 4-6.5 x 2.5-5 cm, ovate to broadly ovate,
asymmetric, base cordate, lobes not or rarely overlapping, apex acuminate, margin
shallowly to moderately lobed, ciliate with a fringe of hairs; adaxial surface green to
reddish green, densely covered with white pilose hairs, abaxial surface red, glabrous;
venation palmate, red, primary veins 5-6, adaxially green to brownish, abaxially
pale red. Inflorescence a simple dichasial cyme, few-flowered, axillary, protandrous,
bisexual; peduncle 3-6 cm long, pink, glabrous; bracts minute, sub-orbicular, margin
slightly fimbriate, deciduous. Male flowers: pedicels 15-18 mm long; tepals 4, pink,
glabrous, two outer tepals ovate to suborbicular, 8-14 x 8-11 mm, apex rounded, two
inner tepals narrowly obovate, 5-12 x 4-6 mm, tip slightly obtuse, androecium yellow,
symmetric, globose; stamens c. 40, filaments fused at base into short column c. 1 mm
long, anthers c. 0.5 mm long, obovate, dehiscing through lateral slits more than half
the length of the anther, apex slightly retuse. Female flowers: pedicels 2.5^ mm long,
bracteoles present, with hairs, ovate, c. 2 mm long, persistent; tepals 4, pink, glabrous,
two outer tepals suborbicular, c. 9.5-12 x 7.5-12.5 mm; glabrous, two inner tepals
elliptic, c. 8-10 x 3.5-5 mm; ovary 3^ x 4 mm (excluding wings), ellipsoid, glabrous,
greenish white, locules 3, placentation axile, wings 3, equal or sub equal, reddish green
triangular, widest point at the middle of the ovary, c. 1 .5 mm long; stigma 3, U-shaped,
stigmatic surface twisted. Fruit: pedicel 2-5 mm long, capsule ovoid, 4-5 x 5-6.5 mm
(excluding wings), dehiscent, splitting along the wing attachments, wing shape as for

Begonia bimaensis, a new species from Indonesia

97

Fig. 1. Begonia bimaensis from cultivated material in Bali Botanie Garden. A-B. Habit; C.
Leaf, adaxial surfaee; D. Leaf, abaxial surfaee; E. Inflorescence; F. Female flower; G. Female
flower; H. Male flower; I. Bracts; J. Ovary transverse seetion; K. Tubers; Scale bars: 1 cm.
(Photos: Gede Wawan Setiadi)

98

Card. Bull. Singapore 67(1) 2015

the ovary, wings widest subapically, 8 mm. Seeds numerous, brown, widely ellipsoid
or ellipsoid, c. 0.35 mm long, 0.2 wide.

Distribution. Endemic to Mt Muria, Bima district, Sumbawa, West Nusa Tenggara, Indonesia.

Habitat. This species grows on steep rocky cliffs in shade, at c. 525 m altitude.

Preliminaty lUCN conservation assessment. Begonia bimaensis is probably a narrowly
endemic species, restricted to Mt Muria which is not fonnally protected. As the species is
known only from a single population with a small number of individuals, a provisonal lUCN
category of CR D is appropriate (lUCN, 2012).

Notes. The epithet 'bimaensis' refers to the Bima district from where the type material
was collected. Begonia bimaensis is unusual in Begonia section Reichenheimia in
Indonesia on account of its tuberous habit, which is similar to Begonia sendangensis, a
species recently described from the neighbouring island of Lombok ( Ardi et. al, 2014).
However, apart from the presence of tubers, the two species are morphologically
dissimilar and Begonia bimaensis can easily be distinguished by the dense white long
hairs on the upper surface of the leaves. The number of tepals in the female flower
and the shape of the ovary wings provide further differences; The female flowers of
Begonia bimaensis have four tepals whereas the female flowers of B. sendangensis
have three tepals; in B. bimaensis, the shape of the wings in the ovaiy is very distinctive,
being triangular and mostly cuneate at apex (versus rounded at base and truncate at the
apex), reddish.

ACKNOWLEDGEMENTS. The authors would like to express their gratitude to Nyoman
Sudiatna for collecting the living specimen (during a fieldtrip supported by the Research
Incentive for Researchers and Engineers No. 053/m/Kp/II/2010 to AK), and to the curators
of the Herbarium of Bali Botanic Garden, BO, E, K, L and SING for allowing us access to
herbarium material. We also thank Mark Hughes (Royal Botanic Garden Edinburgh) for editing
and correcting the English and Wisnu Ardi (Bogor Botanic Gardens) for his assistance.

References

Ardi, W.H. (2014). Two new species of Begonia (Begoniaceae) from Sulawesi, Indonesia.
Edinburgh J. Bot, 71: 259-268.

Ardi, WH. & Hughes, M. (2010). Begonia droopiae Ardi (Begoniaceae). A new species of
Begonia from West Sumatra. Card. Bull. Singapore 62: 1 7-22.

Ardi, W.H., Ardhaka, I.M., Hughes, M., Undaharta, N.K.E., Ginnansyah, D. & Hidayat, S.
(201 3). Two new species of Begonia (Begoniaceae) from Bali and Lombok. Card. Bull.
Singapore 65: 135-142.

Begonia bimaensis, a new species from Indonesia

99

Ardi, W.H., Kusuma, Y.W.C., Lewis, C.E., Risna, R.R., Wiiiadinata, H., Abdo, M.E. & Thomas,
D.C. (2014). Studies on Begonia (Begoniaceae) of the Molucca Islands I: Two new
species from Halmahera, Indonesia, and updated description of Begonia holosericea.
Reinwardtia 14: 19-26.

Lin, C.-W. & Peng, C.-I. (2014). Begonia natunaensis (Sect. Reichenheimia, Begoniaceae), a
new species from Natuna Island, Indonesia. Taiwania 59: 368-373.

Doorenbos, J., Sosef, M.S.M. & de Wilde, J.J.F.E. (1998). The sections of Begonia including
descriptions, keys and species lists (Studies mBegoniacomW). Agric. Univ. Wageningen
Pap. 98: 1-266.

Girmansyah, D. (2009). A taxonomic study of Bali and Lombok Begonia {BegoniacoeiQ).
Reinwardtia 12: 419-434.

Ginnansyah, D. (2012). Begonia ranaiensis (Begoniaceae), a new species from Mt Ranai,
Natuna Island, Indonesia. Kew Bull. 68: 1^.

Girmansyah, D., Wiriadinata, H. & Thomas, D.C. (2009). Two new species and one new
subspecies of Begonia (Begoniaceae) from Southeast Sulawesi, Sulawesi, Indonesia.
Reinwardtia 13: 69-74.

Hughes, M. (2008). An Annotated Checklist of Southeast Asian Begonia. Edinburgh: Royal
Botanic Garden Edinburgh.

Hughes, M. & Pullan, M. (2007). Southeast Asian Begonia Database. Royal Botanic Garden
Edinburgh, http ://elmer,rbge.org.uk/5ego«/n!/ (accessed on 18 Aug. 2014).

Hughes, M., Girmansyah, D., Ardi, W.H. & Nurainas (2009). Seven new species of Begonia
from Sumatra. Card. Bull. Singapore 61: 29-44.

lUCN (20 1 2). lUCN Red List Categories and Criteria: Version 3. 1 . Second Edition. Switzerland,
Gland and UK, Cambridge: lUCN.

Thomas, D.C. & Hughes, M. (2008). Begonia varipeltata (Begoniaceae): Anew peltate species
from Sulawesi, Indonesia. Edinburgh J. Bot. 65: 369-374.

Thomas, D.C., Ardi, W.H. & Hughes, M. (2011). Nine new species of Begonia (Begoniaceae)
from South and West Sulawesi, Indonesia. Edinbutgh J. Bot. 68: 225-255.

Thomas, D.C, Ardi, W.H., Hartutiningsili & Hughes, M. (2009). Two new species of Begonia
(Begoniaceae) from South Sulawesi, Indonesia. Edinburgh J. Bot. 66: 229-238.

Wiriadinata, H. (2011). A new species of Begonia (Begoniaceae) from Sagea Lagoon, Weda
Bay, Halmahera Island, North Moluccas Indonesia. Reinwardtia 13: 263-270.

Gardens’ Bulletin Singapore 67(1): 101-106. 2015
doi: 10.3850/S2382581215000113

101

A new species of Paraboea (Gesneriaceae)

from Thailand

C. Puglisi^, S. Suddee^, R Triboun^ & DJ. Middleton^

'Royal Botanic Garden Edinburgh, 20A Inverleith Row,
Edinburgh EH3 SLR, Scotland, U.K.
c.puglisi@rbge.ac.uk

^The Forest Herbarium, Department of National Parks,
Wildlife and Plant Conservation,

Bangkok, 1 0900, Thailand

^Thailand Institute of Scientific and Technological Research,
Technopolis, Klong 5, Klong Luang,

Pathumthani 12120, Thailand

"'Herbarium, Singapore Botanic Gardens, National Parks Board,
1 Cluny Road, Singapore 259569

ABSTRACT. A new species, Paraboea maculata C.Puglisi, is described. It is compared to
similar species and the habitat on granite rocks, unusual for the genus, is highlighted.

Keywords. Gesneriaceae, Paraboea, Thailand

Introduction

A Paraboea (C.B. Clarke) Ridl. specimen, collected in fruit in Kliao Khitchakut
National Park in Chanthaburi, Thailand, in 2012, was recognised at the time as likely
to be an undescribed species but was not published due to lack of flowering material.
It has now flowered in the Royal Botanic Garden Edinburgh and is here described as
a species new to science.

Paraboea was revised by Xu et al. (2008) but since then the genus has been
considerably enlarged through the inclusion of Trisepalum C.B. Clarke and Phylloboea
Benth. (Puglisi et al., 2011) and by the description of many more species (Chen et al.,
2008; Kiew, 2010, 2012; Chen et al., 2012; Triboun & Middleton, 2012; Xu et al.,
2012; Triboun, 2013; Wen et al., 2013). There are now over 130 species in the genus
with many species being very locally endemic (Xu et al., 2008) which brings with it
conservation concerns (Xu et al., 2008; Triboun & Middleton, 2012). The centre of
diversity of the genus is in Thailand with over 75 species but it is likely that there are
many undiscovered species in neighbouring countries.

Paraboea species are lithophytes, mostly on limestone substrates. A small
number of species are recorded from other substrates, such as Paraboea elegans
(Ridl.) B.L.Burtt (southern Thailand, Peninsular Malaysia) and P. gratiiticola Z.R.Xu
(southern central Vietnam) from granite, and P. pubicorolla Z.R.Xu & B.L.Burtt and

102

Card. Bull. Singapore 67(1) 2015

Paraboea lavandulodora Triboun from sandstone. In Paraboea these are a small
number of exceptions to the otherwise overwhelming preference for limestone. In
Thailand this results in a very uneven distribution of the large number of species,
with very few in the relatively limestone-poor areas of southeastern and northeastern
Thailand.

In Xu et al. (2008) only one collection of 3 . Paraboea species was reported from
the southeastern province of Chanthaburi. No further collections from Chanthaburi
were reported by Triboun & Middleton (2012) and Triboun (2013). Therefore, when a
Paraboea species was collected from granite rocks in Kliao Khitchakut National Park
in Chanthaburi in 2012, it was only the second known collection of any species of the
genus from the province and only the third collection from the whole southeastern
region (the third collection being from the island of Ko Chang). The Khao Khitchakut
plant has a temiinal inflorescence with leaf-like bracts towards the base of the peduncle,
characters which place the species in the Paraboea martinii group (Xu et ah, 2008).
When only known from the fruiting collection, it could already be determined that the
Kliao Kliitchakut plant was not one of the described species in that alliance nor one of
the other species recorded from the southeastern region by Xu et al. (2008). With the
flowering material now available this is confirmed and the species is here described.

Paraboea maculata C.Puglisi, sp. nov.

Differs from other Paraboea species in the Paraboea martinii group, i.e. those with
opposite leaves, campanulate corolla, twisted fruit and a terminal inflorescence, by the
combination of a predominantly white and laterally compressed corolla, the purple
spots at the base of the tube, the sticky glandular secretion on the bracts and calyx,
and the winged petioles. - TYPE: Thailand, Chanthaburi, Khao Khitchakut, Khao
Khitchakut National Park, Kliao Phra Bhat, 1 2°50' 1 4"N 1 02° 1 0'3 "E, 900 m, 27 August
2012, fr., Middleton, D.J., Karaket, P, Suddee, S. & Triboun, P 5675 (holotype E;
isotypes BK, BKF). (Fig. 1)

Lithophytic, caulescent herb to 60 cm high. Stem short and erect, with an indumentum
of sessile and stalked glands, and sparse multicellular, eglandular hairs. Leaves
opposite, congested; petioles winged, 2-9 cm long, those of a pair fonning a distinct
auricle across the node; lamina lanceolate to ovate, 8-19 x 4-10 cm, about twice as
long as wide, surface rugose, apex acute, base rounded and then shortly attenuate
onto the wing of the petiole, margin irregularly crenate, indumentum arachnoid and
deciduous above, with white glands and a pale, thin, interwoven, semi-deciduous
indumentum below, secondai*y veins 10-13 pairs, tertiary veins reticulate, venation
prominent below. Inflorescence a terminal panicle, 25-30 cm long, occasionally
flanked by subtenninal cymes, with at least 4 orders of branching, axes deep red-purple
to bright brown, appearing chestnut brown in herbarium specimens, with a diffuse
arachnoid indumentum or glabrous, bearing a series of sessile, leaf-like, opposite and
decussate pairs of bracts of a progressively decreasing size, with intemodes 5-15 cm
long; uppermost bracts 0.4-1 cm long, 0.3-0. 6 cm wide at the base, sessile, joined at

A new Paraboea from Thailand

103

Fig. 1. Paraboea maculata C.Puglisi. A. Inflorescence. B. Winged petiole. C. Corolla, front
view. D. Corolla and calyx, side view. (Photos: A, C, D: Lynsey Wilson; B: Sadie Barber)

104

Card. Bull. Singapore 67(1) 2015

the base or nearly so, deltoid, concave at the base, proximally red or brown, turning
green towards the tip, densely covered in sessile glands and sticky on both sides but
particularly so on the outer, which is glossy due to an exudate; pedicels 1-1,6 cm long,
glabrous. Calyx with lobes divided to the base, resembling terminal bracts in colour,
indumentum and glossiness; lobes 3-5 mm long, c. 1 mm wide, linear to nan'owly
lanceolate, apex broadly acute. Corolla strongly zygomorphic, campanulate, white
with dark purple-red markings inside towards the base of the tube and around the
staminode insertions, visible from the outer side, laterally and above as a pink shade;
tube laterally compressed with the exception of two lateral bulges miming along its
length, ventrally with 3 narrower bulges, separated by two depressions, running to the
base of the central lower lobe; covered in minute glandular hairs outside making the
corolla sticky but without the glossy appearance of the bracts and calyx; tube 9-1 3 mm
long, slightly oblique, mouth compressed; upper lobes 4-5 x 9-1 1 mm, elliptic; lateral
lobes of the lower lip elliptic, apex obtuse, 5-8 x 5-6 mm; central lobe 6-10 x 4-9 mm
(4 mm at the base widening to 9 mm), slightly folded around the central of the three
bulges running along the tube, apex obtuse, margin undulate. Stamens arising from
the base of the corolla; filaments 0.8-1 cm long, purple both proximally and distally,
with a white to bright yellow knee in the middle, glabrous in the purple areas, densely
covered in glandular hairs on the knee; anthers coherent, 1-2 x 3-3.5 mm, white, with
the connectives and tips of the thecae tinged with purple; thecae strongly divergent;
staminodes 3, white, the lateral ones 1-1.5 imn long, arising 1.5-2 mm above the base
of the tube, with scattered glandular hairs along the filament, the central staminodes c.
0.5 mm long, arising c. 2.5 mm above the base of the tube, glabrous. Disc annular, pale
yellow, c. 0.5 mm long. Gynoecium 11-12 mm long, glabrous; ovary 5-7 mm long,
indistinctly developing into a c. 5 mm long style; style enantiostylous, apically bent,
stigma reduced. Fruit (immature) a strongly twisted capsule, glabrous, green, 2^ cm
long, 1-1.5 mm diameter. Seeds numerous, ellipsoid, slightly compressed, 0.5-0. 7 x
0.15-0.2 mm.

Distribution. Only known from the type locality

Ecology. Evergreen forest, on granite bedrock in deep shade.

Etymology. The epithet refers to the purple markings inside the corolla tube.

Provisional lUCN conservation assessment. Data Deficient (DD). This species is
currently only known from one locality within a National Park. Although there is
some disturbance due to tourism at this one site, it is uncertain how extensive these
possible threats are. There is less than 400 km- of suitable habitat (forest of over 800
m altitude) in the immediate vicinity of the collection locality, which would suggest
an assessment of Endangered based on a restricted Extent of Occurrence if there were
corresponding threats. Similar forest types occur in neighbouring parts of Cambodia,
where the threat levels are even less well known. Even if these forests are included,
the EOO remains less than 2000 km^, still within the boundaries of Endangered based

A new Paraboea from Thailand

105

on EOO. However, a clearer assessment of the populations, the distribution and the
threats would first have to be made before an accurate assessment could be proposed.

Additional specimens examined. THAILAND: Chantanaburi: Khao Khitchakut, Kliao
Khitchakut National Park, Khao Phra Bhat, Middleton, D.J. et al. 5675, cultivated as RBGE
20121417, vouchered 25 Jun 2014 as Puglisi, C. CP2 506 14/1 (E); ibidem, vouchered 15 July
2014 as Atkins, H.J. 18 (SING).

Aote.?.This new species is unusual in being one of the very few species in the genus
to occur on a substrate other than limestone. It quite clearly belongs in the Paraboea
martinii group (Xu et al., 2008), which is characterised by opposite and decussate
leaves, tenninal inflorescence, reduced leaf-like bracts at the base of the peduncle, a
large campanulate corolla, distinct limb, filaments with a bearded knee, and twisted
capsules. Other species in this group include Paraboea doitungensis Triboun &
D.J.Middleton, P. glutinosa (Hand.-Mazz.) K.Y.Pan, P. paramartinii Z.R.Xu &
B.L.Burtt and P. thorelii (Pellegr.) B.L.Burtt. It differs from Paraboea doitungensis
in the shorter calyx lobes, the flower colour, the laterally compressed corolla opening,
the lateral swellings, and the longer fruit; it differs from Paraboea glutinosa in the
broader leaves and the laterally compressed corolla opening; it differs from Paraboea
martinii in the winged petiole, the lack of a matted or glandular indumentum on the
floral axis and the compressed corolla; it differs fi-om Paraboea paramartinii in the
shorter petioles, the broader petiole wings and the more deeply crenate leaf margin;
and it differs from Paraboea thorelii in the broader leaves, lack of glandular hairs on
the inflorescence axes, laterally compressed corolla opening, calyx indumentum, and
the pubescence on the upper leaf surface. We have also compared it to the material
from Cambodia and Vietnam identified provisionally as Paraboea cf. glutinosa by Xu
et al. (2008). It is not the same species as Poilane 28717 from Cambodia, which has
a different leaf indumentum, smaller upper bracts and calyx (but which is otherwise
very similar to Paraboea maculata in shape and stickiness), very much larger lower
bracts, and longer inflorescence intemodes. It is also not the same species as Poilane
16562 from Vietnam, which is an altogether hairier plant than Paraboea maculata,
with bigger bracts and no visible glands.

ACKNOWLEDGEMENTS. We thank the Director and staff of Khao Khitchakut National Park
for the opportunity to study the plants of the Park; Gunnar Ovstebo and Sadie Barber of the
Royal Botanic Garden Edinburgh for their subsequent care and cultivation of the plants; and
Lynsey Wilson and Sadie Barber for the photos.

References

Chen, W.H., Moller, M., Shui, Y.M. & Zhang, M.D. (2008). A new species of Paraboea
(Gesneriaceae) from a karst cave in Guangxi, China, and observations on variations in
flower and inflorescence architecture. Bot. J. Linn. Soc. 158: 681-688.

106

Card. Bull. Singapore 67(1) 2015

Chen, W.H., Moller, M., Zhang, M.D. & Shui, Y.M. (2012). Paraboea hekouensis and R
manhoaensis, two new species of Gesneriaceae from Ch\m..Ann. Bot. Fenn. 49; 179-
187.

Kiew, R. (2010). Two new species of Paraboea (Gesneriaceae) from Peninsular Malaysia and
Thailand. Edinburgh J. Bot. 67: 209-217.

Kiew, R. (2012). Paraboea (Gesneriaceae) in Sabah, Borneo, including a new Malayan

Nat. J. 64; 1-8.

Puglisi, C., Middleton, D.J., Triboun, P. & M5ller, M. (2011). New insights into the relationships
between Paraboea, Trisepalum and Phylloboea (Gesneriaceae) and their taxonomic
consequences. Taxon 60: 1693-1702.

Triboun, P. (2013). Paraboea middletonii (Gesneriaceae), a new species from Thailand. Thai
For. Bull (Bot.) A\: 45-41 .

Triboun, P. & Middleton, D.J. (2012). Twenty new species of Paraboea (Gesneriaceae) from
Thailand. Card. Bull. Singapore 64: 333-370.

Wen, F., Hong, X., Chen, L.Y., Zhou, S.B. & Wei, Y.G. (2013). A new species of Paraboea
(Gesneriaceae) from a karst limestone hill in southwestern Guangdong, China.
Phytotaxa 131; 1-8.

Xu, W.B., Huang, Y.S., Wei, G.F., Tan, W.N. & Liu, Y. (2012). Paraboea angustifolia
(Gesneriaceae): a new species from limestone areas in northern Guangxi, China.
Phytotaxa 62: 39-43.

Xu, Z., Burtt, B.L., Skog, L.E. & Middleton, D.J. (2008). Arevision ofParaboea (Gesneriaceae).
Edinbuj^h J. Bot. 65: 161-347.

Gardens’ Bulletin Singapore 67(1): 107-122. 2015
doi: 10.3850/S2382581215000125

107

New orchid records for Myanmar, including
the first record of the genus Stereosandra

H. KurzweiP & Saw Lwin^

^Herbarium, Singapore Botanic Gardens, National Parks Board,
1 Cluny Road, Singapore 259569
h ubert_kurz\veil @nparks . gov. sg
^Myanmar Floriculturist Association, Yangon, Myanmar
myanmarorchid@gmail.com

ABSTRACT. Nine orchid species, BulhophyUum capnophyton J.J.Verm. et al., Dendrobium
macrostachyum Lindk, D. praecinctiim Rchb.T, Habenaria reniformis (D.Don) Hook.f.,
Liparis distcms C,B. Clarke, L. resupinata Ridk, Micropera thailandica (Seidenf. & Smitinand)
Garay, Oberonia obcordata Lindl. and Stereosandra javanica Blume, are reported as new for
Myanmar. Of particular interest is the Stereosandra as it is the first record of this genus in the
country.

Keywords. Myanmar, Orchidaceae, new distribution records

Introduction

The orchid flora of Myanmar is very rich but as yet poorly known (a brief overview
of the current state of orchid floristics in the country is given in Ormerod & Sathish
Kumar, 2003, 2008), and a complete inventory of all species of Orchidaceae found in
the country is not yet available. Our ongoing studies resulted in nine additions to the
orchid flora of Myanmar, namely BulhophyUum capnophyton J.J.Verm. et al. (syn.
Trias nana Seidenf.), Dendrobium macrostachyum Lindl., D. praecinctiim Rchb.f.,
Habenaria reniformis (D.Don) Hook.f., Liparis distans C.B. Clarke, L. resupinata
Ridk, Micropera thailandica (Seidenf. & Smitinand) Garay, Oberonia obcordata
Lindl. and Stereosandra javanica Blume, all of which are new records for Myanmar
of species found in adjacent countries. Stereosandra is recorded from Myanmar for
the first time.

Brief descriptions, various notes and illustrations are provided for all species.
Detailed locality data below the level of townships are not given here, in order not to
reveal localities which may endanger their populations through over-collecting. The
morphological descriptions are partly based on the specimens from Myanmar and
have been partly extracted from previously published descriptions.

Enumeration of new records

1 . Bulbophyllum capnophyton J.J.Verm., Schuit. & de Vogel, Phytotaxa 166: 110
(2014). - Trias nana Seidenf., Bot. Tidsskr. 71: 24 (1976); Seidenf., Opera Bot. 89:

108

Card. Bull. Singapore 67(1) 2015

164, fig. 106 (1986). - TYPE: Peninsular Thailand, Nam Yong, Seidenfaden, G. &
Smitinand, T GT 6096 (holotype C). (Fig. lA)

Bulbophyllum sp., Seidenf. & Smitinand, Orch. Thailand (Prelim. List): 388, fig. 294
(1961). Based on A.F.G. Kerr 0361 (BK) from Thailand.

Herb to 2 cm tall, glabrous. Pseudobulbs spaced along a creeping rhizome at intervals
of 10-14 mm, globular, somewhat flattened, 5-8 mm high, 6-10 mm in diameter,
1 -leaved. sessile, fleshy, duplicate, oblong-lanceolate, acute, 10-21 mm long.
Inflorescences 1 -flowered, scape 1-2 mm long, pedicel and ovary c. 4 mm long,
bract 1-2 mm long, hyaline, cup-shaped. Flowers light yellow, the lip deeper yellow.
Sepals spreading, oblong-elliptic, acute, 5-veined, 7-8 x 2.3-3 mm. Petals narrowly
ovate, acute, 1 -veined, 2.5-3 x c. 1.5 mm; lip oblong, obtuse, 2. 3-2. 8 x c. 1 mm,
with forward curved auricles at the base; column c. 1 .7 mm long (excluding its anther
process), stelids insignificant, anther process fleshy, oblong, somewhat spoon-shaped,
c. 1.6 mm long, apex emarginate.

Distribution. Peninsular regions of Thailand and Myanmar.

Ecology and phenology. Very little habitat infomiation on this species is available.
The specimen from Myanmar was growing as an epiphyte at an elevation of about 400
m. In neighbouring Thailand the specimen Kerr 0361 (BK) was collected in savanna
(Seidenfaden & Smitinand, 1961 : 388). In the distribution area of the species flowering
has been recorded in November and January.

Material examined. MYANMAR. Taninthayi Region: Yebyu Township, 22 Nov 2012, Saw
Lwin, Pan Khet Khet &. law Oo Wai TNRO 162 (SING, SING [spirit], herbarium of Taninthayi
Nature Reseive Education Centre).

Notes. The specimen from Myanmar, Lwin et al. TNRO 162, matches Seidenfaden’s
type very well in its stixicture (Seidenfaden, 1976a). However, it differs in the slightly
larger size of the leaf and flowers: leaf 20-21 mm long (as opposed to 10-14 mm in the
type); sepal c. 8 mm long (as opposed to c. 7 mm in the type); petal c. 3 mm long (as
opposed to 2. 5-2,8 mm in the type); lip c. 2.8 mm long (as opposed to 2. 3-2. 5 mm in
the type). The epithet nanum was not available in Bulbophyllum when Trias nana was
moved into it due to Bulbophyllum nanum De Wild.

2. Dendrobium macrostachyum Lindl., Gen. Sp. Orchid. PI. 78 (1830). - Callista
macrostachya (Lindl.) Kuntze, Revis. Gen. PI. 2: 655 (1891). - TYPE: Sri Lanka,
1829, Macrae, J. 77 (holotype K-LINDL! [K000364613]). (Fig. 2A)

Dendrobium stuartii F.M.Bailey, Proc. Roy. Soc. Queensland 1: 12 (1884);
Vaddhanaphuti, Wild Orch. Thailand, ed. 4, 125, inch colour photo (2005); Seidenf. &

New records of Myanmar orchids

109

Fig. 1. A. Anther of Bulbophylliim capnophyton JJ.Verm., Schuit. & de Vogel. From Lwin
et al TNRO 162, B. Column of Micropera thailandica (Seidenf & Smitinand) Garay. From
Lwin et al. TNRO 61. C. Inflorescence of Ohemnia obcordata Lindl. Tmm Kurzv^feil & Twin
KL 2664. D. Lip of Oberonia obcordata Lindl. From Kurzweil & twin KL 2664. E. Habit of
Stereosandra javanica Blume. from Twin et al. TNRO 5. F. Flower of Stereosandra javanica
Blume. From Lwin et al. TNRO 5. G. Lip of Stereosandra javanica Blume. From Lwin et al.
TNRO 5. Scale bars: A-D, F-G = 1 mm, E = 20 mm. (Drawn by Joshua Yang)

no

Card. Bull Singapore 67(1) 2015

Fig. 2. A. Dendrobium macrostachyum Lindl. Vvom Lwin MPO 050. B. Dendrobium praecinctum
Rchb.f. ¥rom Lwin SL 49. C. Habenaria reniformis (D. Don) Hook.f. Vvom Lwin et al. MPO 020.
D-E. Liparis distans C.B. Clarke. From Kurzweil & Lwin KL 2460 (Photos: Saw Lwin)

New records of Myanmar orchids

111

JJ.Wood, Orchids Penins. Malaysia Singapore 389, fig. 175, pi. 26d (1992). - Callista
stuartii (F.M.Bailey) Kiintze, Revis. Gen. PL 2: 655 (1891). TYPE: Australia, drawing
by R.D.Fitzgerald in Austral Orch. 2(3): t. 6 (1888) (neotype designated by Clements,
1989).

Dendrohium tetrodon auct. non Rchb.f. ex Lindl.: Seidenf., Opera Bot. 83: 34, fig. 13
(1985).

Herb with slender pseudobulbous and sometimes almost wiry, pendulous stems to
70 cm long, glabrous except for the lip. Stems slightly zig-zagged, intemodes 2-2.5
cm long, entirely covered by white membranous sheaths, leafy throughout. Leaves
distichous, sessile, spreading, ovate to oblong-lanceolate, acute or acuminate, 4-8 x
0.9-2. 5 cm. Inflorescences to 2.5 cm long, with l-3(^) flowers; bracts oblong-ovate,
acute, 2. 5-2. 6 x 1 .5-1 .8 mm. Flowers 20-25 mm across, sepals and petals spreading
or recurved; whitish to greenish yellow, lip with red-brown or purple veins on the
side lobes, ageing to yellow; median sepal oblong-lanceolate, acute, 13-15 x 3,5^
mm; lateral sepals similar but often slightly longer; mentum c. 6 mm long, narrowly
conical; petals elliptic-lanceolate, acute or obtuse, as wide as the sepals, 13-1 5 x 3.5^
mm; lip quadrately-ovate and very obscurely 3-lobed, 17-18 x c. 12 mm, unspurred,
hairy particularly on the margin and midrib, basal part inrolled around the column,
callus in the basal lip portion of three diverging ridges; column 3.5^ mm long.

Distribution. Widespread in tropical Asia from Sri Lanka, India and the Himalayas to
eastern Malesia; also found in northern Australia.

Ecology and phenology. The specimen from Myanmar was growing as an epiphyte
on medium-sized road-side trees at an elevation of 1150 m. Elsewhere the species is
found in wet and dry low-altitude and montane forest to 1220 m (Jayaweera, 1981;
Seidenfaden & Wood, 1992). In the distribution area of the species flowering has been
recorded in January and February.

Material examined. MYANMAR. Mandalay Region: Kyaukpadaung Township, Jan 2010,
Saw Lwin MPO 050 (SING, SING [spirit]). Very uncommon in this locality, growing in a small
population of about 20 individuals.

Notes. A widespread species with rather small flowers in bunches of mostly 1-3 on
leafless stem. The taxonomy of this species was previously disputed (Christenson
& Wood, 2003), in particular its relationship to Dendrohium aphyilum (Roxb.)
C.E.C.Fisch., but this has now been clarified (Schuiteman, 2011).

3. Dendrohium praecinctum Rchb.f., Gard. Chron., n.s., 7: 750 (1877); N.Pearce &
RJ.Cribb, FI. Bhutan 3(3): 416, pi. 23 [top right] (2002); Jin et al., Acta Bot. Yunnan.
32: 332 (2010). - Callista praecincta (Rchb.f.) Kuntze, Revis. Gen. PI. 2: 655 (1891).
TYPE: Unlocalised, cult. Veitch s.n. (holotype W). (Fig. 2B)

112

Card. Bull. Singapore 67(1) 2015

Dendrobium pauciflorum King & Pantl., J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 64: 332
(1895); Seidenf., Opera Bot. 83: 97, fig. 57, pi. XIIc {19S5), - Dendrobium sikkimense
A.D.Hawkes & A.H.Heller, Lloydia20: 124 (1957), nom. illeg. -TYPE: India, Sikkim,
July 1895, Pantling, R. 172 (holotype K! [K000943939]).

Herb, pendent stems to 70 cm long, but often much shorter. Stems leafy in the distal
half, branching from nodes in the upper part of the stem, intemodes 1 .4-4 cm long.
Leaves distichous, to 12, lanceolate or linear-lanceolate, acute, sessile, 3-9 x 0.7-1. 2
cm. Inflorescences axillary, arising from apical nodes of the leafless stem, with 1-5
flowers; floral bracts ovate, obtuse, c. 2 mm long. Flowers to 20 mm long, not opening
widely; yellow to whitish yellow, sepals and petals with prominent purple marginal
areas, midlobe and side lobes of lip with red spots; pedicel and ovary 5-10 mm long;
median sepal lanceolate or lanceolate-ovate, obtuse, 3-10 x up to 4 mm; lateral
sepals ovate, weakly falcate, keeled, 4-10 x 2-4 mm; petals lanceolate-ovate, usually
ciliolate, 4-10 mm long and up to 3 mm wide; lip rather deeply 3-lobed, somewhat
clawed, 4-12 x 3-8 mm, hispid-fimbriate on the margins; lateral lobes narrow, tooth-
like; mid-lobe oblong, flat; disk with three prominent, thickened veins; column broad,
with a foot 2-7 mm long.

Distribution. Northeastern India and eastern Himalayas to Myanmar, Thailand and
southwestern China (Yunnan).

Ecology and phenology. Habitat information of the specimen from Myanmar has not
been recorded. In Sikkim and Bhutan the plants are epiphytic in evergreen broad-
leaved forest at elevations ranging from 1000 to 1330 m, and rarely up to 1850 m
(Pearce & Cribb, 2002). In the distribution area of the species flowering has been
recorded in June and July.

Material examined. MYANMAR. Shan State: Ho Pone Township, 2011, Saw Lwin SL 49
(herbarium of the Myanmar Floriculturist Association).

Notes. Easily recognised by the prominent purple edges on the pale yellow flowers
and the rather deeply 3-lobed lip which is strongly hairy to fringed on the margins.
As far as we are aware no vouchered record of the occurrence of Dendrobium
praecintum in Myanmar has previously been made. However, a photograph of an
unidentified Dendrobium species from Putao in northern Kachin State was published
by Nyan Tun (2014: 254) and was later refeiTed to this species by Paul Onnerod (pers.
comm.). Interestingly, the disnibutioii ‘Myamnar’ was also listed in a publication
newly reporting Dendrobium praecinctum in Yunnan, China (Jin et al., 2010), but no
specimen from Myanmar was cited and therefore this listing appears to be based on a
sight-record only.

New records of Myanmar orchids

113

4. Habenaria reniformis (D.Don) Hook.f., FI, Brit. India 6: 152 (1890); Seidenf.,
Dansk Bot. Ark. 31(3): 132, fig. 85 (1977); S.C.Chen & P.J.Cribb in Z.Y.Wu et al.
(eds), FI. China 25: 149 (2009); Kurzweil, FI. Thailand 12(1): 133, fig. 76 (2011). -
Listera reniformis D.Don, Prodr. FI. Nepal.: 28 (1825). - Neottia reniformis (D.Don)
Spreng., Syst. Veg. 3: 707 (1826). - TYPE: Nepal, 1821, Wallich, N. 7067 (lectotype
K-W!, designated by Seidenfaden (1977)). (Fig. 2C)

Habenaria clovisii Gagnep., Bull. Soc. Bot. France 78: 68 (1931). -TYPE: Cambodia,
Stung-streng, 1866-1868, Thorel, C. 2143 (syntypes P! [P00439699, P00439700]);
Vietnam, near Dalat, 17 Oct 1924, Evrard, F 1480 (syntype P! [P00439698]).

Herb to 20 cm tall, glabrous. Tubers globose to oblong, to 1 cm in diameter, fleshy.
Stems slender, wiry, to 1 mm in diameter. Leaves 2-4, basal and adpressed to the
substrate, orbicular, elliptic or ovate, acute or obtuse, 1.4-4 x 1-2.7 cm. Inflorescences
lax, with 3-7(-10) flowers; with or without a spreading amplexicaul leafy bract near
the base; sterile bracts 4-6(-9), mostly erect, lanceolate, acuminate, 0.5-1. 8 cm long;
rachis 3-8 cm long; floral bracts lanceolate or ovate-lanceolate, acuminate, 7-10 x

1.5- 2 mm, much shorter than the ovary. Flowers 6.5-9 mm in diameter, resupinate;
green, greenish-white, yellow-green or brownish; ovary twisted mamly in its basal
portion, cylindric-fusiform, indistinguishable from the pedicel, 13-18 mm long;
median sepal erect, concave, ovate or ovate-oblong, obtuse, 3-veined, 3.5-4(-5) x

1.5- 3 mm; lateral sepals spreading or reflexed, obliquely ovate-lanceolate, subacute
or acute, 3-veined, 3.5-4(-6) x 1.7-3. 4 mm; petals fonning a hood with the median
sepal, falcately elliptic-lanceolate or linear-triangular, subacute or obtuse, 1 -veined,
3^.5 X 0.8-1. 6 mm, sometimes widened at the base; lip 4.5-10 mm long, either
unlobed and linear, or 3-lobed with filifonn side lobes to 29 x 0. 1-0.3 mm; spur either
present or absent, if present saccate or cylindric and to 7.6 mm long; column 1-2.3
mm long; anther canals insignificant; stigmas subcylindric or clavate, 1 .5-2 mm long.
Capsule fusiform-elliptic, 9.5-10 x 2.2-3 mm.

Distribution. India and Nepal to Indochina and southern China; also recorded from
northern Sumatra.

Ecology and phenology. No ecological information on the specimens from Myanmar is
available except that the species grows terrestrially. Elsewhere found in grassy places
in forests, in bamboo thicket and scrubland at elevations ranging from 1 50 to 825 m,
sometimes also in cracks of limestone rocks (Chen et al., 2009; Kurzweil, 2011). In
the distribution area of the species flowering has been recorded in August, October and
November.

Material examined. MYANMAR. Mandalay Region: Kyaukpadaung Township, 26 Oct 2009,
Saw Lwin, Pan KhetKhet & Sandar Maung MPO 020 (SING). Growing in a small population.

114

Card. Bull. Singapore 67(1) 2015

Notes. Habenaria reniformis is characterised by small greenish or brownish flowers
with sepals mostly less than 4.5 mm long. The lip shape is variable, ranging from entire
to 3-lobed (Banerji & Pradhan, 1984: 36; Chen et al., 2010: 193; Baretto et al, 2011:
260), and the spur can be absent or cylindrieal and up to 7.6 mm long (Seidenfaden,
1977: 133; Kurzweil, 2011: 133; Baretto et al., 2011: 260). The rather robust anther
and the long protruding stigmas are characteristic of the species (Seidenfaden, 1977).

In the collection from Myanmar, Lwin et al. MPO 020, the petals are basally
widened to more than double their width above. The petals and the lip are basally
united with the column. The lips of the two plants in this collection are mostly unlobed.
The only exception is one flower which has an asymmetric 3-lobed lip and appears to
be abnonually developed: one side lobe is linear-filiforai and 4 mm long, and the other
one semicircular and only 0.5 mm long.

Habenaria reniformis is very close to H. hnmidicola Rolfe and H. poilanei
Gagnep., both of which differ by having larger leaves, consistently deeply 3-lobed lips
and longer spurs. The Habenaria diphylla (Nimmo) Dalzell group is also very close.
The relationships among these species are not fully resolved and it is not yet clear if all
of these taxa can be mamtained as separate species. As already noted by Seidenfaden
(1977), further research is needed to reach a final position on their status.

5. Liparis distans C.B. Clarke, J. Linn. Soc., Bot. 25: 71 (1889); Seidenf , Dansk Bot.
Ark. 31(1): 76, fig. 50 (1976); N.Pearce & P.J.Cribb, FI. Bhutan 3(3): 212 (2002);
S.C.Chen et al. in Z.Y.Wu et al. (eds), FI. China 25: 225 (2009). - Leptorkis distans
(C.B.Clarke) Kuntze, Revis. Gen. PL 2: 671 (1891). - Stichorkis distans (C.B. Clarke)
Marg., Szlach. & Kulak, Acta Soc. Bot. Poloniae 77: 38 (2008). - TYPE: India,
Nagaland, Kohima, 21 Oct 1885, Clarke, C.B. 41071 (syntype K! [K000387820],
CAL); Kohima, Clarke, C.B. 41099 (syntype K, not found); Kohima, 21 Oct 1885,
Clarke, C.B. 41105 (syntype K! [K000387819]); Kohima, 21 Oct 1885, Clarke, C.B.
41574 (syntype K! [K0003 87821]). (Fig. 2D-E)

Liparis macrantha Hook.f, Hooker’s Icon. PI. 19: t. 1854 (1889). - TYPE: India,
Nagaland, Prain, D. 44 (lectotype K, designated by Seidenfaden (1976b)).

Liparis yunnanens is Rolfe, J. Linn. Soc., Bot. 36: 8 (1903). - TYPE: China, Yunnan,
Henry, A. 10485 (holotype K! [K000364665]).

Liparis oxyphylla Schltr., Repert. Spec. Nov. Regni Veg. Beih. 4: 63 (1919). - TYPE:
China, Yunnan, Henry, A. 12593 (lectotype K! [K0003 64666], designated here;
isolectotype E! [E00286229]).

Herb mostly 20-40 cm tall, pseudobulbous, glabrous. Pseudobulbs clustered,
subcylindric or narrowly ovoid, 2-6(-9.5) cm long, slightly compressed. Leaves
2, distichous, shortly and indistinctly petiolate, articulate, spreading, nan'owly
oblanceolate, acute or acuminate, 15-35 x 1-2.8 cm. Inflorescences arcuate or

New records of Myanmar orchids

115

pendulous, 15-39 cm long, lax, with up to 12 distant flowers; peduncle somewhat
compressed, narrowly winged, with 2-3 sterile bracts; floral bracts narrowly triangular,
6-12 mm long. Flowers medium-sized, olive-green to dull yellow; pedicel and ovary
14-22 mm long; sepals narrowly lanceolate, obtuse, mostly 10-16 x 3-5 mm, margins
often revolute; petals linear-filifoiin, obtuse, 10-16 x c. 2 mm; lip ovate, obovate or
elliptic, 10-14 x 10-11 mm, basal part a short claw, margin irregularly denticulate,
rounded or obtuse at the apex, with a 2-lobed callus at the base, unspurred; column
arcuate, 5-6 mm long, narrowly winged in the upper part. Capsule ellipsoid or
narrowly obovoid-oblong, 12-18 x 6-7 mm.

Distribution. Widespread from northeastern India and the eastern Himalayas to
Myanmar, Indochina and southwestern and southern China. Also in the Philippines.

Ecology and phenology. The specimen from Myanmar was growing as an epiph34e in
primary mountain forest. Elsewhere the species is lithophytic or epiphytic in various
forest types at elevations up to 2400 m (Chen et al., 2009). In Vietnam often found in
limestone regions (Averyanov, 2013). In the distribution area of the species flowering
has been recorded between September and February.

Material examined. MYANMAR. Kachin State: Border region of Putao and Nogmung
Townships, 7 Mar 2007 (fruiting at the time of collection; flowering in cultivation), Kurzweil,
H. & Saw Lwin KL 2460 (herbarium of the Myanmar Floriculturist Association).

Notes. Distinct with its large flowers in lax inflorescences and the wide lip with
denticulate margins.

6.LiparisresupinataRid\., J. Linn. Soc., Bot. 22: 290 (1886); Seidenf., DanskBot. Ark.
31(1): 88, fig. 60 (1976); N.Pearce & P.J.Cribb, FI. Bhutan 3(3): 209, pi. 7 [top right]
(2002); S.C.Chen et al. in Z.Y.Wu et al. (eds), FI. China 25: 228 (2009). - Leptorkis
resupinata (Ridl.) Kuntze, Revis, Gen. PI. 2: 671 (1891). - Platystyliparis resupinata
(Ridl.) Marg., Richardiana 7: 39 (2007). - TYPE: India, Darjeeling, 1844, Griffith, W.
s.n. (syntype K-LINDL! [K000873787]); India, Kliasia, sine collector 986 (syntype
K-LINDL! [K000873786]); India, Khasia, Griffith, W s.n. (syntype K-LlNDL!
[K000873788]); India, Sikkim, 5000 ft [1524 m], Hooker, J.D. 90 (syntype K-LINDL!
[K000873785]); India, Kliasia, Lobb, T. 122 (syntype K-LINDL! [K000873784]);
India, Kliasia, Hooker, J.D. & Thompson, T. 90 (syntype K-LINDL! [K000873783]).
(Fig. 3A-B)

Liparis ridleyi Hook.f, Hooker’s Icon. PI. 19: t. 1887 (1889). - Leptorkis ridleyi
(Hook.f) Kuntze, Revis. Gen. PI. 2: 671 (1891). - TYPE: India, Sikkim, 3000-5000
ft [91 4-1 524 ni], Hooker, J.D. 89 (syntype K-LINDL! [K000873782]); India, Sikkim,
Hooker, J.D. s.n. (syntype K-LINDL! [K000387829]); Griffith s collector, Kew Distr.
5081 (syntype K); India, Darjeeling, 5500 ft [1676 m], 11 Mar 1875, Clarke, C.B.

116

Card. Bull Singapore 67(1) 2015

Fig. 3. A-B. Liparis resupinata Ridl. From Lasi BawkNaw BW 32. C. Micropera thailandica
(Seidenf. & Smitinand) Garay. FromLwm et al. TNRO 61 (Photos: A, H. Kurzweil; B-C, Saw
Lwin)

27235 (syntype K-LINDL! [K000387827]; India, Darjeeling, 5000 ft, [1524 m], 2 Mar
1876, Clarke, C.B. 27088 (syntype K-LINDL! [K000387828]).

Herb 12-28 cm tall, with densely arranged pseudobulbous stems. Pseudobulbs 1.5-5
X 0.3-0. 6 cm, made up of a few nodes, with leaves in the upper half Leaves 2-A,
papery, subsessile, narrowly oblong to linear-lanceolate, acute to acuminate, 6-13
X 0.4-1. 2 cm, articulate. Inflorescences arching or pendent, lax, 10-50-flowered;
peduncle slightly zig-zagged, 5-8 cm long, with several sterile bracts 0.5-0. 9 cm
long; rachis 8-18 cm long; floral bracts lanceolate, acute, concave, 3-5 x 1-1.5 mm.
Flowers to 2 mm wide and 5 mm high when seen from the front; in the specimen
from Myanmar pale greenish orange-yellow with orange on the basal part of the lip
including its auricles, in other parts of the distribution area reported as pale green,
greenish yellow, golden brownish or creamy; pedicel and ovary 5-7 mm long; sepals
subequal, spreading, oblong or elliptic-oblong, obtuse or acute, 1 -veined, 3-4 x 1-1.8
mm; petals reflexed downwards, narrowly linear, obtuse, 2-3.5 x c. 0.3 mm; lip knee-

New records of Myanmar orchids

117

like, bent with the lower portion erect and parallel to the column and the distal portion
spreading forwards, broadly elliptic-oblong or broadly ovate-oblong, 2.5-3 mm long,
unspun*ed, base auriculate and with a bilobed callus, apex obtuse, margins incurved,
forming a pouch; column erect, 1-2.8 mm long, with two suborbicular wings, each
with a prominent pendulous filifonn appendage. Capsule obovoid-oblong, c. 5 x 2-3
mm.

Distribution. India and Himalayas to Indochina and southwestern China (Xizang and
Yunnan). Also recorded in the Gaoligongshan Mountains in western Yunnan Province
(Jin et ah, 2009), which is very close to where our specimen from Myanmar was
collected.

Ecology and phenology. The specimen from Myanmar was growing as an epiphyte
in degraded mixed broad-leaved forest with much bamboo and moss-covered trees
at an elevation of 2100 m. Elsewhere the species is found in evergreen broad-leaved
and coniferous forest from 1500 to 2500 m (Pearce & Cribb, 2002; Chen et ah, 2009;
Averyanov, 2013). In the distribution area of the species flowering has been recorded
from October to March.

Material examined. MYANMAR. Kachin State: Waingmaw Township, 10 Mar 2011, Stephen
Lasi Bawk Naw BW 32 (herbarium of the Myamnar Floriculturist Association). Locally
connnon.

Notes. One of the epiphytic Liparis species with few-leaved pseudobulbs. Well-
characterised by the shape of the small flowers that have a broadly oblong lip with
an obtuse and pouched apex and the column with characteristic pendulous filiform
appendages on the wings.

7. Micropera thailandica (Seidenf. & Smitinand) Garay, Bot. Mus. Leafl. 23: 187
(1972); Seidenf., Opera Bot. 95: 126, fig. 72 (1988); Vaddhanaphuti, Wild Orch.
Thailand, ed. 4, 186, inch colour photo (2005). - Camarotis thailandica Seidenf &
Smitinand, Orch. Thailand (Prelim. List) 712, fig. 529 (1965). - TYPE: Thailand,
Satun Province, Feb 1961, Seidenfaden, G.& Smitinand, T. GT 4118 (holotype C).
(Fig. IB, 3C)

Herb with monopodial growth. Stems to 30 cm long, glabrous, covered by the dark,
rugose bases of the leaves. Leaves about 1 0, distichous, spaced at intervals of about
2 cm, flat, conduplicate, oblong-linear, apex unequally bilobed with rounded lobes,
8-15 X 1-1,5 cm, fleshy. Inflorescences 17-23 cm long, several-flowered; peduncle
10-15 cm long, with 2-3 sheathing sterile bracts 0. 3-0.4 cm long; rachis 8-10 cm
long, unbranched; bracts broadly triangular, obtuse, 2-2.5 x c. 4 mm, fleshy. Flowers
yellow or light orange-yellow, lip base sometimes white, sepals sometimes tipped with
brown, column white or cream; median sepal elliptic-ovate, apex broadly rounded.

118

Card. Bull. Singapore 67(1) 2015

to 9 X 3.5 mm, concave, fleshy; lateral sepals similar but somewhat shorter and wider,
adnate to the column-foot; petals elliptic-lanceolate, acute, to 8 x 3.3 mm; lip 7 mm
long, deeply boat-shaped with a conical spur 4-5 mm long in its distal part, obscurely
3-lobed, very fleshy; midlobe triangular, c. 2 x 2-2.5 mm; side lobes inconspicuous,
broadly triangular, edges minutely serrate; callus a triangular tongue at the base of
the midlobe, split at the end, below this a rounded callus at the backwall of the spur
(terminology after Seidenfaden, 1988); spur with a longitudinal septum; column c. 6
mm long, venti'ally with a sudden horizontal edge with two short lateral horns below
the stigma, above which the column narrows to about one half of its width below,
higher up widening again under the anther and the rostellum, rostellum beak c. 2 mm
long, not or only slightly twisted.

Distribution. Myanmar, Thailand and Vietnam.

Ecology and phenology. The specimen from Myanmar was found in primary evergreen
forest at about 220 m, growing as an epiphyte on tall trees. In Thailand recorded at
elevations of between 700 and 1000 m (Seidenfaden, 1988). In the distribution area of
the species flowering has been recorded in February and March.

Material examined. MYANMAR. Taniiithayi Region: Yebyu Township, 712 ft [217 m], 25
Mar 2012, Saw Lwin, Pan Khet Khet & Zaw Oo Wai TNRO 61 (SING, SING [spirit], herbarium
of the Myanmar Floriculturist Association). Locally common.

Notes. This species is unmistakable in its lip and column structure. It is known from
several collections in Thailand and is also found in Vietnam; in addition it is here
newly reported for Myanmar. Seidenfaden & Smitinand (1965) compared it with the
Malaysian Camarotis adnata (Ridl.) Holttum (now included in Micropera fuscolutea
(Lindl.) Garay), which is superficially similar, but this species differs in details of its
lip and column (Seidenfaden & Wood, 1992).

8 . Oberonia obcordata Lindl., Fol. Orchid. 8: 7 (1859); Seidenf, Dansk Bot. Ark.
33(1): 30, fig. 19 (1978); N.Pearce & P.J.Cribb, FI. Bhutan 3(3): 230 (2002); S.C.Chen
et al. in Z.Y.Wu et al. (eds), FI. China 25: 240 (2009). - Malaxis obcordata (Lindl.)
Rchb.f. in W.G.Walpers, Ann. Bot. Syst. 6: 216 (1861). - Iridorkis obcordata
(Lindl.) Kuntze, Revis. Gen. PL 2: 669 (1891). - TYPE: India, Sikkim, Hooker, J.D.
& Thompson, T. 112 (syntype K-LINDL! [K000974237]); India, Darjeeling, 1844,
Griffith, W. s.n. (syntype K-LINDL! [K000974238]); India, Kliasia, Griffith, W. s.n.
(syntype K-LINDL! [K000974231]). (Fig. IC-D, 4A)

Oberonia treutleri Hook.f, Hooker’s Icon. PI. 18: t. 1786 (1888). - Iridorkis treutleri
(Hook.f.) Kuntze, Revis. Gen. PI. 2: 669 ( 1 89 1 ). - TYPE: India, Sikldm, 6000 ft [ 1 829
m], 1 Dec 1874, Treutler, W.J. 1151 (holotypeK! [K000387726]).

New records of Myanmar orchids

119

Fig. 4. A. Fruiting specimens of Oberonia obcordata Lindl. From Lasi Bawk Naw BW 8. B.
Fruiting specimen of Stereosandra javanica Blume in its natural habitat in southern Myanmar.
From Lwin et al. TNRO 5 (Photos: A, H. Kurzweil; B, Saw Lwin)

Herb to 9 cm tall, with leafy stems to 4 cm tall, glabrous. Leaves (3-)5-8, distichous,
equitant, bilaterally eompressed, fleshy, linear-oblong, slightly falcate, acute to
aeuminate, mostly 1.5-5 x 0. 5-0.9 cm, base not articulate. Inflorescences terminal,
densely or subdensely many-flowered, 4. 5-7. 5 cm long; peduncle connate to lower
part of uppermost leaf and therefore appearing to arise from its blade, with few to
many erect sterile braets about 0.5 cm long; rachis slender, 1 .5-6 em long; floral braets
verticillate, lanceolate or ovate-lanceolate, apex acuminate-caudate, 2-A mm long,
much longer than the flowers, margin slightly erose. Flowers 1-1.5 mm in diameter;
reddish brown or red; pedicel and ovary 0.5-1. 5 mm long; median sepal ovate, acute,
to 1.1 X 0.6 mm; lateral sepals broadly ovate, obtuse, to 1.1 x l mm; petals linear-
oblong, subaeute, to 0.9 x 0.5 mm; lip 3-lobed, not spurred, to 1.6 mm long; side
lobes divergent, ovate-oblong, obtuse, to 0.5 mm long; midlobe obcordate or reniform,
emarginate, to 1 x 0.8 mm, with a prominent basal callus; column to 0.5 mm long.
Capsule ovoid, to 5 x 4 mm.

Distribution. Himalayas, northeastern India, Myanmar and southwestern China to
Thailand.

Ecology and phenology. The specimens from Myanmar were found in forest at various
elevations (see below). Elsewhere either epiphytic or lithophytic, usually at elevations

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Card. Bull. Singapore 67(1) 2015

up to 3000 m (Pearce & Cribb, 2002; Chen et al., 2009). In the distribution area of
the species flowering has been recorded between July and December and in March.
Flowering and fruiting plants can sometimes be found at the same time.

Material examined. MYANMAR. Kachin State: Putao Township, undisturbed evergreen
forest, 1 500 m, 2 1 Mar 2009, Kurzweil, H. & Saw Lwin KL 2664 (SING); Waingmaw Township,
degraded mixed forest, 2200-2300 m, 9 Mar 2011, Stephen Lasi Baw’k Naw BW 8 (RAF,
herbarium of the Myanmar Floriculturist Association). Locally common in both localities.

Notes. Very distinct due to the inflorescence which is connate to the uppermost leaf,
the long and whorled floral bracts, and the small flowers with their large lip side lobes
and the basal lip callus.

9. Stereosandra javanica Blume, Mus. Bot. 2: 176 (1856); Seidenf , Dansk Bot. Ark.
32(2): 173, fig. 107 (1978); Hedge & Rao, Indian J. Forest. 10(3): 196 (1987); Seidenf.
& J.J. Wood, Orchids Penins. Malaysia Singapore 143, fig. 60 (1992); N.Pearce &
P.J.Cribb, FI. Bhutan 3(3): 582 (2002); S.C.Chen et al. in Z.Y.Wu et al. (eds), FI. China
25: 207 (2009); H.A.Pedersen, FI. Thailand 12(2): 632, fig. 342 (2014). - TYPE:
Western Java, Bantam Province, Kuhl, H. & van Hasselt, J.C.A. s.n. (? holotype L!).
(Fig. lE-G, 4B)

Stereosandra pendula Ki'aenzL, Bot. Tidsskr. 24: 11 (1901).- TYPE: Thailand, South-
Eastern, Ko Chang, Schmidt, J. s.n. (not known to exist; see Seidenfaden, 1978: 173).

Herb terrestrial, holomycotrophic, 15-40 cm tall, glabrous. Corm ellipsoid or ovoid,
1. 5-4.4 X 0.6-2.4 cm. Stems leaf-less, slender to robust, 1^ mm in diameter; with 2-7
erect and sheathing sterile bracts 1.1-1. 8 cm long. Inflorescences laxly 3- to many-
flowered; rachis 3-19 cm long; floral bracts lanceolate, acuminate, 3.5-11.6 x 1. 2-4.3
mm, mostly shorter than the ovary. Flowers resupinate; white or cream with purple
markings and purple-tipped sepals, petals and lip; pedicel and ovary distinct from
each other, ovary ellipsoid, 3-8.2 mm long, pedicel 1.5-5 mm long; sepals lanceolate,
acute to acuminate, 8-10.1 x 1. 6-2.2 mm; petals similar, linear-lanceolate, acute to
acuminate, 7-9.2 x 2.2-2. 6 mm; lip ovate-lanceolate, obtuse to subacuminate, entire to
irregularly undulate-crenate, 5-8 x 2. 3-3. 5 mm, concave; with two ellipsoid wart-like
calli at the base; column incurved, 3^.2 mm long. Capsule ellipsoid to subspherical,
5-13 X 3-7 mm.

Distribution. Widespread from the eastern Himalayas through the whole of tropical
and subtropical Asia to New Guinea and the islands of the southwestern Pacific Ocean
(east to Samoa),

Ecology and phenology. The specimens from Myanmar were found in bamboo-rich
hill evergreen forest near a stream at an elevation of 1 80 m. Elsewhere the species is

New records of Myanmar orchids

121

found in primary evergreen forest from 100 to 1500 m (Seidenfaden & Wood, 1992;
Chen et aL, 2009; Pedersen, 2014). In the distribution area of the species flowering
occurs mostly between March and June but has also been recorded between December
and February. At the locality in southern Myanmar both flowering and fruiting plants
were seen at the same time.

Material examined. MYANMAR. Taninthayi Region: Yebyu Township, 23 Mar 2012, Saw
Lwin, Pan KhetKhet & Zaw Oo Wai TNRO 5 (SING, SING [spirit]). Locally common.

Notes. This is a new generic record for the country. Given its overall distribution
(tropical Asia from Thailand and Peninsular Malaysia eastwards to New Guinea;
Pridgeon et al., 2005 : 528) the occurrence of Stereosandra in Myanmar is not surprising.
In neighbouring Thailand Stereosandra javanica is also found in areas adjacent to the
border with Myanmar. Known localities in southern and southwestern Thailand are the
provinces of Ranong and Phetchaburi, both bordering Myanmar’s Taninthayi Region
where our specimen was collected. We suggest that Stereosandra javanica may be
more widespread in the southern part of Myanmar, but has probably been overlooked
because of its inconspicuous appearance.

ACKNOWLEDGEMENTS. Fieldwork on Mount Popa, in Kachin State, and in Taninthayi
Nature Reserve was made possible by the Myanmar Forest Department and we would like to
thank the officials concerned for providing the necessary pennits. We would like to acknowledge
Htoo Foundation for providing research funding for the orchid surveys on Mount Popa. Thanks
are also due to Stephen Lasi Bawk Naw, Brang Aung and local foresters for their assistance in
the field. We would like to thank Paul Onnerod for advice on the newly reported species, Jaap
J. Vermeulen for providing infonuation on BuJbophyllum capnophyton, and Jonathan Gregson
for checking herbarium material at BM. Thanks are also due to .loshua Yang for making the line
drawing, and the fu'st author would like to acknowledge the National Parks Board in Singapore
for finaneial support.

References

Averyanov, L. (2013). The orchids of Vietnam illustrated survey. Part 4. Turczaninowia 16(1):
5-163.

Baneiji, M.L. & Pradhan, P. (1984). The Orchids of Nepal Himalaya. Lieehtenstein, Vaduz: J.

Cramer, A.R. Gantner Verlag Kommanditgesellschaft.

Baretto, G., Cribb, P.J. & Gale, S. (2011). The Wild Orchids of Hong Kong. Malaysia, Kota
Kinabalu: Natural Histoiy Publications (Borneo) & China, Hong Kong: Kadoorie Farm
and Botanic Garden.

Chen, S.-C., Liu, Z.J., Zhu, G.H., Lang, K.-Y, Tsi, Z.-H., Luo, Y.B., Jin, X.H., Cribb, P.J.,
Wood, J.J., Gale, S.W., Ormerod, P., Veniieulen, J.J., Wood, H.P., Clayton, D. & Bell,
A. (2009). Orchidaceae. In: Wu, Z.Y., Raven, P.H. & Hong, D.Y. (eds) Flora of China,
vol. 25: 1-570. China, Beijing: Science Press & USA, St. Louis: Missouri Botanical
Garden Press.

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Chen, S.-C., Liu, Z.J., Zhu, G.H., Lang, K.-Y., Tsi, Z.-H., Luo, Y.B., Jin, X.H., Cribb, RJ.,
Wood, JJ., Gale, S.W., Ormerod, R, Vermeulen, J.J., Wood, H.R, Clayton, D. & Bell,
A. (2010). Orchidaceae. In: Wu, Z.Y, Raven, RH. & Hong, D.Y (eds) Flora of China,
vol. 25, illustrations: 1-666. China, Beijing: Science Rress & USA, St. Louis: Missouri
Botanical Garden Rress.

Christenson, E. & Wood, H. (2003). The mystery of Limodorum aphyllum - Resolving lingering
taxonomic problems concerning two dendrobiums. Orchids (West Palm Beach) 72:
774-777.

Clements, M.A. (1989). Catalogue of Australian Orchidaceae. Austral. Orchid Res. 1: 1-160.
Jayaweera, D.M.A. (1981). Orchidaceae. In: Dassanayake, M.D. & Fosberg, F.R. (eds) A
revised Handbook to the Flora of Ceylon, vol. 2. India, New Delhi: Amerind Rublishing
Co.

Jin, X.-H., Zhao, X.-D. & Shi, X.-C. (2009). Native Orchids from Gaoligongshan Mountains,
China. China, Beijing: Science Rress.

Jin, X.-H., Dai, Z.-Q., Liu, Q.-Y. & Ju, X.-Y. (2010). Miscellaneous taxonomic notes on
Orchidaceae from China. Acta Bot. Yunnan. 32: 331-333.

Kurzweil, H. (2011). Habenaria. In: Santisuk, T. & Larsen, K. (eds) Flora of Thailand, vol.
12(1): 79-149. Thailand, Bangkok: The Forest Herbarium, Department of National
Parks, Wildlife and Riant Conservation.

Nyan Tun (2014). Wild Orchids of Myanmar. Myamnar, Yangon: Daw Tin Tin Aye (Green Leaf
Publishing House).

Ormerod, P. & Sathish Kumar, C. (2003). Orchidaceous additions to the flora of Burma
(Myanmar). Rheedea 13: 43-50.

Ormerod, P. & Sathish Kumar, C. (2008). Orchidaceous additions to the flora of Myanmar 2.
Rheedea 18: 75-80.

Pearce, N.R. & Cribb, RJ. (2002). Flora of Bhutan: Including a record of plants from Sikkim
and Darjeeling, vol. 3(3), The Orchids of Bhutan. UK: Royal Botanic Garden Edinburgh
& Bhutan: Royal Government of Bhutan.

Redersen, H.^. (2014). Stereosandra. In: Santisuk, T. & Balslev, H. (eds) Flora of Thailand,
vol. 12(2), pp. 631-633. Thailand, Bangkok: The Forest Herbarium, Department of
National Parks, Wildlife and Plant Conservation.

Pridgeon, A.M., Cribb, P.J., Chase, M.W. & Rasmussen, F.N. (2005). Genera Orchidacearum,
vol. 4 (Epidendroideae, part 1). UK: Oxford University Press.

Schuiteman, A. (2011). The strange case of Dendrobium aphyllum. Orchid Rev. 119: 104-110.
Seidenfaden, G. (1976a). Conhibutions to the orchid flora of Thailand VII. Bot. Tidsski: 71:
1-30.

Seidenfaden, G. (1976b). Orchid genera in Thailand IV. Liparis L.C.Rich. Dansk Bot. Ark.
31(1): 1-105.

Seidenfaden, G. (1977). Orchid genera in Thailand V. Orchidoideae. Dansk Bot. Ark. 31(3):
1-149.

Seidenfaden, G. (1978). Orchid genera in Thailand VI. Neottioideae Lindl. Dansk Bot. Ark.
32(2): 1-195.

Seidenfaden, G. (1988). Orchid genera in Thailand XIV. Fifty-nine vandoid genera. Opera Bot.
95: 1-398.

Seidenfaden, G. & Smitinand, T. (1961). The Orchids of Thailand - A preliminary list. Part III.
Thailand, Bangkok: The Siam Society.

Seidenfaden, G. & Smitinand, T. (1965). The Orchids of Thailand -A preliminary list. Part IV
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Denmark, Fredensborg: Olsen & Olsen.

Gardens’ Bulletin Singapore 67(1): 123-127. 2015
doi: 10.3850/S2382581215000137

123

A new species of Zingiber (Zingiberaceae)

from Lao P.D.R.

M.R Newman

Royal Botanic Garden Edinburgh, 20A Inverleith Row,
Edinburgh EH3 5LR, Scotland
m.newman@rbge.ac.uk

ABSTRACT. Zingiber nitens M.F.Newman from Lao RD.R. is described and illustrated.
Keywords. lUCN conservation assessment, Lao RD.R., new species, Zingiber

Introduction

Rhizomes of a wild ginger collected in 2011 by Dr Vichith Lamxay of the National
University of Laos and his colleagues were donated to the Royal Botanic Garden
Edinburgh for cultivation. As the material they collected was without flowers but had
the remains of a temiinal inflorescence, the rhizomes were labelled Alpinia. When
the plant flowered in the living collection at Edinburgh in 2012, it was immediately
apparent that this was a new species of Zingiber.

Zingiber Mill, is among the larger genera of Zingiberaceae with 144 species
(Govaerts et ah, 2015) distributed from Sri Lanka and India to southern China, Japan
and SE Asia. Schumann (1904) classified the genus into four sections, two of which,
Zingiber section Dymczewiczia (Horan.) Benth. and Z. section Pleuranthesis Benth.,
contained species with terminal inflorescences. What Schumann did not observe was
that some species in Zingiber section Zingiber and Z. section Ciyptanthiiim Horan.,
such as Zingiber barbatum Wall., Z. gramineum Noronha ex Blume, and Z. junceum
Gagnep., could produce inflorescences both radically and tenuinally (Triboun et al.,
2014). Schumann (1904) placed Zingiber barbatum in Zingiber section Cryptanthium
Horan, and Z. gramineum in Zingiber section Zingiber (which he called Zingiber
section Lampugium Horan., nom. inval.); Gagnepain (1906) placed Z. junceum in
Zingiber section Zingiber (again as Zingiber section Lampugium).

Now, there are at least eight species of Zingiber known to produce terminal
inflorescences. They are found from the eastern Himalaya (Z. capitatum Roxb. and
Z. clarkei King ex Baker in India and Sikkim) to Papua New Guinea (Z. brevifolium
K.Schum.). Zingiber confine Miq. is from southern China, Z. pellitum and Z.
rufopilosum Gagnep. are found in Continental SE Asia, Z. gramineum is Irom Java,
and Z. marginatum Roxb. is a poorly known species of unknown origin,

Theerakulpisut et al. (2012) estimated the phylogeny of 23 species of Zingiber
using ITS sequences and concluded that Zingiber sections Zingiber, Dymczewiczia
and Pleuranthesis were individually weakly supported but together formed a well-
supported clade sister to a well-supported clade consisting of Z. section Cryptanthium.

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Card. Bull. Singapore 67(1) 2015

The following description and illustration were made from a full-grown, living
plant and the lUCN assessment uses the criteria given in lUCN version 3.1 (2012).

Zingiber nitens M.F.Newman, sp. nov.

Belongs to Zingiber section Dymczewiczia (Horan.) Benth. because the inflorescences
are produced tenninally on the leafy shoots; similar to Zingiber capitatum Roxb. in
its slender habit with narrow leaves and tenninal inflorescences but differs from it by
its glabrous, glossy, dark green bracts wliich remain green even at fruiting (not green
bracts which turn red at fruiting, and are sparsely to densely villose at the margins). -
TYPE: Originally a living collection from Lao P.D.R., Bolildiamxai prov., Khamkeut
district. Ban Thongpe, Nakai-Nam Theun NPA, Lao- Vietnam Border Protected Area,
18°1L54" N 104°35'52.9"E, 573 m altitude, primary evergreen forest along river,
1 August 2011, Lamxay, K, Lanorsavanh, S., Souvannakoummai, K, & Somphone
VL2188, grown on as cultivated material at RBGE acc. no. 201 11 043 A, vouchered
and selected as type as Newman, M.F. 2647 (holotype L, inch spirit). (Fig 1, 2)

Clump-forming herb 0.65-1 m tall. Rhizome c. 1 cm in diameter. Leafy shoots
composed of c. 12 leaves, leaf sheaths dark brownish green, especially lower ones,
coarsely white villose, hairs pointing in all directions; pseudostem c. 1 cm in diameter;
ligule 3-5 mm long with truncate apex, translucent green when young, soon becoming
light brown, especially at margin, coarsely white pubescent; swollen petiole light
green. Leaf blade narrowly elliptic, 18-25 x 2-3 cm, glabrous and shiny adaxially,
sparsely white villose abaxially, especially on midrib, base rounded, apex very long
acute. Inflorescence a terminal, erect thyi'se, 9-13 x 1.3-2 cm, cylindrical to fusifonn,
composed of 9-1 5 bracts, wrapped round and obscuring rhachis. Bracts subtending to
3-4 flowers, somewhat obovate, c. 35 x 33 mm, glossy, dark green, glabrous, surface
with minute oil glands, apex acute, margin translucent. First flower of cincinnus
ebracteolate, subsequent flowers with a boat-shaped bracteole, open to base, c. 20
X 6.5 mm, translucent at base, green at apex, glabrous. Flower exserted from its
bract; 4 cm long, calyx 13-16 x c. 4 mm, tubular, inflated, translucent, glabrous,
split halfway down one side, apex with 2 short, blunt teeth; floral tube 30-3 1 mm
long, widenmg slightly towards apex, white at base, pale yellow at apex, lobes pale
yellow, glabrous, dorsal corolla lobe triangular, 19-21 x c. 6 mm, margins slightly
inrolled, lateral corolla lobes 18-20 x 5-5.5 mm; lateral staminodes triangular, 2.5-6
mm long, almost free from labellum, pale yellow, sometimes with a few red dots;
labellum elliptic, c. 18 x 1 1-12.5 mm, dark maroon with small yellow dots and yellow
patch in throat, margins deflexed, apex bifid for 3 mm. Stamen: filament 2.5-3 x 3
mm long, yellow; anther c. 1 1 x 4 mm, connective tissue yellow, thecae dehiscing by
longitudinal slits; anther crest 9-10 mm long, wrapped around stigma, curved down
into cleft of lip, dark maroon. Style white, glabrous, stigma white, round, scarcely
wider than style, ostiole with ring of straight cilia; epigynous glands 2, subulate, c. 3.6
mm long; ovary cylindrical, c. 3.5 x 2.5 mm, glabrous, incompletely trilocular at base,
the three placentas forming a very short axis bearing 10-14 ovules, unilocular above.

New Zingiber from Lao P.D.R.

125

Fig. 1. Zingibernitens M.F.Newman. A. Habit. B. Ligule. C. Inflorescence. D. Cincinnus, bract
removed. E. Bract. F. Bracteoles, showing size range. G. Calyx. H. Floral tube, corolla lobes,
labellum and lateral staminodes, abaxial and adaxial views. I. Stamen, style, stigma and dorsal
corolla lobe. J. Detail showing position of style in corolla tube. K. Upper style and stigma,
adaxial and lateral views. L. Ovary (cross-sectioned in apical part). Scale bars: A= 10 cm, B,
J— L = 5 mm, C— I = 2 cm. Drawn by Claire Banks from RBGE living accession 201 1 1043 A.

126

Card. Bull Singapore 67(1) 2015

Fig. 2. Zingiber nitens M.F.Newman, from RBGE living accession 20 11 1043 A. (Photo: Mark
Newman)

placentas petering out on inside walls not far from base. Infmctescence c. 10 cm long,
mature fruits unknown.

Phenology. The wild collection, Lamxay et al. VL2188, had finished flowering in
August but detailed observations on phenology will require further collections.

Distribution & ecology. Zingiber nitens is only known from the type locality, where it
grows in primary evergreen forest along a river at c. 570 m altitude.

Provisional lUCN conservation assessment. Data Deficient (DD). Zingiber nitens
is a perennial, terrestrial herb which may be expected to support a certain amount
of disturbance. The area in which it was found has a degree of legal protection and
there is no evidence that the species is being harvested or otherwise targeted. Until the
extent of the wild population is better known, it is impossible to give a more precise
assessment.

New Zingiber from Lao P.D.R.

127

Etymology. The epithet “nitens”, Latin for shining, refers to the shiny braets of the
inflorescence.

Notes. One of the vegetative characters of Zingiber, and many other Zingiberoideae,
is the presence of an abscission layer at the base of the pseudostem which allows the
plant to shed its pseudostems for the duration of the dormant season, whether that is
the dry season in the monsoon tropics, or the winter in more northerly areas. Zingiber
nitens possesses an abscission layer but, in cultivation in Edinburgh, it does not shed
all its pseudostems. Instead, the laminae become rather chlorotic looking in the winter
but persist. It is not known what happens in wild populations.

All parts of the living plant give off a foetid smell when bruised, and the flowers also
smell foetid.

ACKNOWLDEGEMENTS. Claire Banks is gratefully acknowledged for the line drawing
used in Fig. 1.

References

Gagnepain, F. (1906). Zingiberacees nouvelles de I’herbier du Museum. Bull. Soc. Bot. France
53: 132-150.

lUCN (2012). lUCN Red List Categories and Criteria: v. 3.1. 2nd ed. Switzerland, Gland &
UK, Cambridge: lUCN.

Schumann, K.M. (1904). Zingiberaceae. In: Engler, H.G.A. (ed) Das Pflanzenreich: regni
vegetabilis conspectus, IV, 46, Heft 20. Leipzig: W. Engelmann.

Theerakulpisut, P., Triboun, P., Mahakham, W., Maensri, D., Khampila, J. & Chantaranothai, P.
(2012). Phylogeny of the genus Zingiber (Zingiberaceae) based on nuclear ITS sequence
data. Kew Bull. 67(3) 389-395.

Triboun, P., Larsen, K. & Chantaranothai, P. (2014). Akey to the genus Zingiber (Zingiberaceae)
in Thailand with descriptions of 1 0 new taxa. Thai J. Bot. 6(1): 53-77.

Govaerts, R., Newman, M. & Lock, J.M. (2015). World Checklist of Zingiberaceae. Royal
Botanic Gardens, Kew. http://apps.kew.org/wcsp/ (accessed on 11 Mar. 2015).

Gardens’ Bulletin Singapore 67(1): 129-142. 2015
doi: 10.3850/S2382581215000149

129

Taxonomic studies on Zingiber (Zingiberaceae) in China I:
Zingiber kerrii and the synonymy of Z. menghaiense

and Z. stipitatum

L. J. Leong-Skoraickova^ & N.H. Xia^

^Key Laboratory of Plant Resources Conservation & Sustainable Utilization,
South China Botanical Garden, Chinese Academy of Sciences,
Guangzhou, 510650 People’s Republic of China
nhxia@scib.ac.cn

"University of Chinese Academy of Sciences,

Beijing, 100049 People’s Republic of China
bailin@scib.ac.cn

^Herbarium, Singapore Botanic Gardens, National Parks Board,

1 Cluny Road, 259569 Singapore
j ana_skomicko va@nparks .gov. sg

ABSTRACT. A general introduction to Zingiber is provided, along with an outline of the
materials and methods used in this paper and which are to be used in our subsequent taxonomic
studies on Zingiber in China. Based on an investigation of the protologues, and on comparisons
of both the type specimens and living materials collected from the type localities and their
immediate vicinities. Zingiber stipitatum S.Q.Tong and Z. menghaiense S.Q.Tong are here
reduced to synonymy under Z. kerrii Craib. A previous lectotypification of Zingiber kerrii is
discussed and amended. The existence of mixed collections among the isotypes of Zingiber
stipitatum is highlighted and a proposal to exclude the vegetative parts belonging to Z.
neotruncatum from the type material is made. A description and a colour plate of Zingiber
kerrii based on a collection from China are provided.

Keywords. India, Myanmar, synonym, Thailand

Introduction

Zingiber Mill. (Zingiberaceae, Zingibereae) is an economically important genus, best
known for the ginger of commerce. Zingiber officwaJe (L.) Roscoe. The genus is
widely distributed in tropical to warm-temperate Asia (Wu & Larsen, 2000). According
to The Plant List (2013), 244 names have been published in this genus, corresponding
approximately to 100-150 species (Theilade, 1999; Wu & Larsen, 2000; Kishor &
Leong-Skomickova, 2013). Strongly supported as monophyletic (Kress et al., 2002),
this genus is easily recognised among Zingiberaceae either by the flower structure (the
horn-shaped anther crest embracing the upper part of the style) or by the vegetative
character (the existence of a swollen part of the petiole, widely known as a pulvinus).

The currently accepted infrageneric classification of Zingiber recognises four
sections, based on the nature and position of the inflorescence: (1) sect. Zingiber,

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Card. Bull. Singapore 67(1) 2015

having a spike on an erect and, usually, long peduncle; (2) sect. Cryptanthium Horan.,
characterised by radical inflorescences composed of a spike appearing at ground level
with a, usually, short procumbent peduncle; (3) sect. Pleuranthesis Benth., with spikes
breaking though the leaf sheaths laterally; (4) sect. Dymczewiczia Benth., with tenuinal
inflorescences. A recent molecular study based on a single marker and limited material
indicates that Zingiber sect. Dymczewiczia and Zingiber sect. Pleuranthesis are not
well segregated from Zingiber sect. Zingiber (Theerakulpisut et ah, 2012). As more
studies involving broader sampling are certainly needed before a new classification
can be formally proposed, the traditional sectional treatment is utilised in our current
work.

The last comprehensive taxonomic revisions of the genus are more than a
century old (Horaninow, 1862; Schumann, 1904), covering only 23 and 55 species
respectively. Several regional studies have been made in S & SE Asia in the meantime,
e.g. Ridley (1909), Valeton (1918), Loesener (1930), Holttum(1950), Smith (1988a,b),
Theilade (1996), Theilade (1999), Sabu (2003), Sabu (2006) and Triboun (2006).

Poor or missing type material, difficulties in the preservation of important
floral characters on herbarium specimens through traditional drying methods, as
well as variability of certain morphological characters coupled with hybridisation
and polyploidy in some genera, make Zingiberaceae taxonomically one of the most
challenging plants groups (Larsen, 1980; Theilade, 1999; Skomickova & Sabu,
2005; Leong- Skomickova et al., 2010; Ardiyani et al., 2011). Our current poor
understanding, particularly of the larger ginger genera, is further exacerbated by the
fact that many previous workers limited their area of study to a single country, rather
than working monographically, leading to repeated descriptions of the same taxa. A
number of taxonomists have written that the ideal approach to the taxonomic study
of gingers is to work with living fertile material or material well-preserved in spirit
(Smith, 1988a; Theilade, 1999; Skomickova & Sabu, 2005; Leong-Skomickova et
al., 2010). This may involve revisiting and collecting plants from the type localities
in order to take precise notes aided by good photo-documentation, and to preserve at
least fertile bracts, bracteoles and flowers (or better still the entire inflorescence) in
spirit, including having several flowers preserved separately to avoid damage (Burtt
& Smith, 1976). A thorough review of the pertinent literature of the species across its
phyto-geograph ical range and good communication between taxonomists working on
gingers is also crucial.

The genus Zingiber in China

Fifty-three names have been associated with Zingiber taxa in China, of which 46 are
based on types from China. The most recent taxonomic treatment of Zingiber is that
of Wu & Larsen (2000) in the Flora of China. In this work, 42 species are recognised,
with 34 being endemic to China. The genus is mostly restricted to the subti'opical
zone of China, i.e. south of the Qinling-Huaihe Line (Zhang & Tang, 1991). While
the account by Wu & Larsen (2000) serves as a useful overview, the work was based

Zingiber kerrii and the synon3Tny of Z. menghaiense and Z. stipitatum

131

mostly on the study of imperfectly preserved herbarium material, of which most
are not accompanied by spirit material. This resulted in short and often incomplete
descriptions lacking basic rhizome and floral characters, and in doubtful synonymies.
About a quarter of Chinese Zingiber species are known only from the type collections
and their identities are poorly understood. There are also a number of presumably
well-known species to which a certain name has been applied for an extended period
of time, sometimes simultaneously to more than one species, and yet an understanding
of what these species really are is not straightforward. These issues can only be
um-avelled with detailed taxonomic work.

While a thorough revision of Zingiber in China is far from complete, accounts
for various regional floras in neighbouring countries are in progress. It is therefore
useful to disseminate our existing results which include clarifications of the identities
of various taxa, the establishment of new synonyms, and the descriptions of new taxa.
The present paper, clarifying the identities of two species from Yunnan, Zingiber
menghaiense S.Q.Tong and Z. stipitatum S.Q.Tong, is the first in the series. It,
therefore, includes a more detailed introduction to the genus as outlined above, as well
as an explanation of the materials and methods applied in our studies, the details of
which will not be repeated in subsequent papers.

Material and methods

Our work on a revision of Zingiber in China was initiated in 201 1 . The protologues of
all published names along with all other pertinent literature on the genus were collated
and reviewed. Searches were made in the relevant herbaria for the original material
or type material identified from the protologues. All Chinese specimens of Zingiber
from China were examined at CDBI, EMA, GXMG, GXMI, HGAS, HITBC (in older
literature often referred as YNTBI), IBK, IBSC, KUN, LBG, PE, SYS. Additional
material from China and neighbouring countries (particularly Thailand, Laos and
Vietnam) was mostly accessed as hi-resolution digital images from the following
herbaria: AAU, BK, BKF, BM, C, E, G, HAST, K, KFRI, L, P, SING, TAI, TAIF, US,
W, WU. Since 2012 the first author has conducted extensive fieldwork to re-collect
fertile material from type localities or their vicinities. Spirit collections, consisting
of fertile bracts, bracteoles and single flowers, were made in the field for further
study. Extensive photographic documentation was also made following the protocols
established by Leong-Skomickova et al. (e.g. 2014a, 2014b). Rhizomes were collected
and brought for planting to the greenhouse in South China Botanic Garden for further
observation. The terminology in general follows Beentje (2010) and the recent works
of Kishor & Leong-Skomickova (2013) and Leong-Skornickova et al. (2014a). While
some previous works (e.g. Theilade, 1999; Wu & Larsen, 2000) treated the labellum as
a tri-lobed stmcture composed of a mid- lobe and two side lobes (implymg the absence
of staminodes in the genus Zingiber), this approach has not been supported by any
moi*phological study and is not in accord with our current knowledge of ginger flower
stmcture. In Zingiberaceae, of the six stamens, only the median posterior stamen

132

Card. Bull. Singapore 67(1) 2015

of the inner whorl is fertile, while the remaining two are sterile and comiate into a
labellum. Of the outer whorl, one of the stamens is fully reduced, while the remaining
two appear as sterile staminodes (ranging in shape from large and petaloid to small
teeth-like stmctures), flanking the stamen or adnate to the labellum (e.g. Larsen et ah,
1998). Petaloid staminodes strongly adnate to labella occur in other Zingiberaceae
genera, e.g. Siliquamomum or Siphonochilus. We therefore agree with the conclusion
of various authors (e.g. Ridley, 1899; Holttum, 1950; Smith, 1988a; Larsen et al.,
1998) that the side lobes in Zingiber are staminodes, which in some species are free to
the base and in others are well developed and might be either fully or partially connate
to the labellum. In a few species they may be almost fully reduced or even missing.
The way we have measured the labellum and lateral staminodes is shown in Fig. 1.
The degree of comiation is also an important character which should be included in
descriptions.

The identities of Zingiber menghaiense and Z. stipitatum

During initial herbarium work it was noticed that the holotypes of Zingiber menghaiense
S.Q.Tong and Z. stipitatum S.Q.Tong appear to belong to the same taxon (Fig. 2 A-C).
At the same time it was also noticed that two of the tliree existing isotypes of Zingiber
stipitatum consisted of mixed collections (Fig. 2C). Furthermore, two specimens
originally identified as Zingiber stipitatum and collected from the vicinity of the type
locality of Z. menghaiense {Pei, S.J. 11344 in KUN^ were redetennined as Z. kerrii
Craib by Dr Pramote Triboun in 2002. This prompted us to investigate the complex of
these three names in greater detail.

Zingiber menghaiense (Tong, 1987) was described from two collections from
Xishuangbanna in south Yunnan, China, Tong, S.Q. & Li, A.M. 32860 from Menghai
Xian and Tong, S.Q. & Li, A.M. 32902, Irom Jinghong city, with the HITBC specimen
of the former collection being designated as the holotype (Fig. 2 A). When first
described, it was inexplicably compared to Zingiber zerumbet (L.) Roscoe ex Sm., a
very different species with incurved bracts, rather than to other more similar species
with appressed bracts.

In the same article, Tong (1987) described Zingiber stipitatum based on a
single collection, Tong, S.Q. & Liao, C.J. 24836 from Ruili Shi, Dehong Zhou, in
the southwest of Yunnan (holotype HITBC48857, Fig. 2B). In the protologue it was
compared to Zingiber menghaiense. Both species have a narrow, lanceolate, glabrous
lamina of similar size and shape, a long erect peduncle, an oblong or narrowly ovoid
spike, pale green bracts with a purple red margin, a whitish labellum, and lateral
staminodes with red patches on both sides of the bases. Zingiber stipitatum was said
to differ by the following characters: (1) the ligules 6 imn long, emarginate (versus 6
mm long, bilobed in Z. menghaiense)', (2) the labellum orbicular with emarginate apex
and petiolate [meaning it has an attenuate base or is clawed] (versus labellum narrowly
obovate, apex bifid in Z. menghaiensey, (3) the entire lateral staminodes which are
shortly acuminate at the apices (versus bifid at apices in Z menghaiense).

Zingiber kerrii and the synonymy of Z. menghaiense and Z. stipitatum

133

Fig. 1. Schematic outlines of selected types of labellum (pink) and lateral staminodes (blue)
oeeurring in the genus Zingiber. From left to right, lateral staminodes nearly free from labellum,
lateral staminodes eonnate to labellum by basal %, lateral staminodes absent, (a) length of
labellum, (b) width of labellum, (e) length of lateral staminodes, (d) width of lateral staminodes.

A comparison of the holotypes of the two species reveals that they are quite
similar to each other. Also, the holotype of Zingiber stipitatum does not match the
protologue as the ligules are c. 3 mm long and clearly 2-lobed. Though shorter, the
ligules of Zingiber stipitatum are also very similar to those of Z. menghaiense in their
nearly coriaceous texture, being slightly pubescent, with a prominent raised vein on
the base and the membranous margin. As pointed out by Triboun (2006), the length of
ligules in Zingiber varies according to the position of leaves on the pseudostem and
those in the middle part of the shoot are usually longer than the others. Three isotypes
of Zingiber stipitatum were located at HITBC and KUN herbaria (HITBC49032,
KUN0833208 and KUN0833209). Further examination of these sheets revealed
that while the isotype at HITBC is of the same taxon as the holotype, the plant parts
mounted on the other two isotypes at KUN (KUN0833208, KUN0833209 (Fig. 2C))
belong to two different species. The ligules on the shoots are emarginate and c. 6 mm
long, consistent with the description and the painting in the protologue, while they
conflict with the protologue and the holotype in the lower lamina surface, sheaths
and ligules being densely villous (they are glabrous in the protologue and holotype).
These two shoots are easy to recognise as Zingiber neotruncatum T.L. Wu, K.Larsen &
Turland, which also occurs in Ruili as mentioned in its protologue (Tong, 1987), while
the inflorescences are certainly of Z. stipitatum (the inflorescence of Z. neotruncatum
has shorter peduncles and is more or less bent and without a dark brown tinge on the
tips of the bracts). As confirmed by the first author in the held. Zingiber neotruncatum
is quite common at the type locality of Z. stipitatum. It appears that the description
of Zingiber stipitatum in the protologue is based on these two different species. The
character of the ligules being emarginate, the only vegetative character used by Tong
(1987) to distinguish Zingiber stipitatum from Z. menghaiense, is derived from the
leafy shoots of Z. neotruncatum present in the two isotypes. After their exclusion, all
the remaining original materials show no critical differences to the type specimens
of Zingiber menghaiense. Further observation of floral characters was made through

134

Card. Bull Singapore 67(1) 2015

Fig. 2. Zingiber kerrii Craib. A. Holotype of Z. menghaiense S.Q.Tong {Tong, S.Q. & Li, AM.
32860, HITBC Acc. No. 048849). B. Holotype of Z. stipitatum S.Q.Tong {Tong, S.Q. & Liao,
C.J. 24836, HITBC Acc. No. 049032). C. One of the two mixed isotypes of Z. stipitatum
S.Q.Tong; the inflorescence belongs to Z. stipitatum, the leafy shoot belongs to Z. neotruncatum
{Tong, S.Q. & C. J. Liao, C.J. 24836, KUN Acc. No. 0833209; barcode 1219333). D. Lectotype
of Zingiber kerrii Craib {Kerr, A.F.G. 1290; barcode K000255235); reproduced with the kind
permission of the Director and the Board of Trustees, Royal Botanic Gardens, Kew.

Zingiber kerrii and the synon3Tny of Z. menghaiense and Z. stipitatum

135

the field work of the first author in 2012 and 2013. More than ten individuals were
found at the type locality of Zingiber menghaiense, with the flowers examined having
entire lateral staminodes (as opposed to staminodes with bifid apices as stated in the
proto logue) but all other characters matched the protologue and the type specimens well.
According to our observation of several other Zingiber species (i.e. Z. longyangjiang
Z.Y.Zhu), the phenomenon of lateral staminodes being bifid to various degrees occurs
occasionally in populations with otherwise entire staminodes, and is therefore not to
be considered a reliable character to distinguish two species. The shape of the labellum
also varies within and between populations and we have observed varying degrees of
narrowing at the bases and notches at the apices which covers the shapes of labella as
depicted in the line drawings of Zingiber stipitatum and Z menghaiense published in
the protologues. At the type locality of Zingiber stipitatum, which is 360 km away (as
the crow flies) from the type locality of Z. menghaiense, only Z. neotruncatum was
found. However, plants matching the description of Zingiber stipitatum were found
within 20 km of the type locality and these are identical to those from the type locality
of Z. menghaiense. It is, therefore, concluded here that Zingiber stipitatum is the same
taxon as Z. menghaiense.

The identity of Zingiber kerrii Craib

Zingiber kerrii Craib was described in 1912 from a collection from Northern Thailand
(Chiang Mai Kerr 1290, Fig. 2D) (Craib, 1912). It was included in the revisions of
Zingiber in Thailand by Theilade (1999) and Triboun (2006) and both works included
photos of the inflorescence and flower. The species was also reported to occur in Shan
state in Myanmar (Kress et al., 2003) and in Manipur state in India (Thongam et al.,
2013). Compared to the Thai collections (Theilade, 1999; Triboun, 2006), the Indian
collection is slightly different by having the labellum and lateral staminodes creamy
white throughout. Our collection of Zingiber menghaiense, approximately 400 km
away from the type locality of Z. kerrii, shows more resemblance to Z. kerrii than the
Indian record, especially in the coloration of the labellum and lateral staminodes. Even
though the laminas are only c. 2.5 cm wide in the type material of Zingiber kerrii,
and described as 2.3 cm wide by Triboun (2006), they can reach up to 5 cm wide in
some other specimens in Thailand, similar to the measurements reported from India
(Thongam et al., 2013). Chinese collections consistently have laminas 3.5^ cm wide.
In conclusion, we agree with Triboun that Zingiber menghaiense (and also Z. stipitatum
as noted above) is the same taxon as Z. kerrii. Therefore, Zingiber menghaiense and Z.
stipitatum are reduced to synonymy under Z. kerrii. Colour plates, including a flower
dissection of Z. kerrii from Yunnan, are provided (Fig. 3, Fig. 4).

Theilade (1999), in her revision, indicated the presence of syntypes of Kerr
1290 at BK, BM, E and K and proposed the lectotypification of Zingiber kerrii with
a specimen deposited at K. However, the Kew herbarium has two specimens of Kerr
1290, both of which contain both a leafy shoot and an inflorescence, and as there
is no indication that these two specimens are part of a single preparation, a further

136

Card. Bull Singapore 67(1) 2015

Fig. 3. Zingiber kerrii Craib. A. Young shoots (inset: leaf sheath and ligule). B. Inflorescences
(left at anthesis, right in fruit). C. Old rhizome with root tubers and section of young rhizome
and root tubers. D. From left: Bract, bracteole, mature capsule (dehisced) and seeds enclosed
in arils. From Bai, L. 13080301 and Bai, L. 12091402, from the type locality of Zingiber
menghaiense. (Photos: L. Bai)

second step lectotypification is needed. In conformity with the ICN (Arts. 8.3, 9.17)
we designate here one of these two specimens, which also contains a flower dissection
(barcode K000255235), as the lectotype.

Zingiber kerrii and the synonymy of Z. menghaiense and Z. stipitatum

137

Fig. 4. Zingiber kerrii Craib. A. Flower (front view). B. Flower (semi-side view). C. Flower
dissection (from left to right: bract, bracteole, single flower in side view, dorsal corolla lobe,
two lateral corolla lobes, labellum with lateral staminodes partially basally connate, stamen
with upper part of style in the groove between two anther thecae. From the type locality of
Zingiber menghaiense, Bai, L. 13080301. (Photos: L. Bai)

Zingiber kerrii Craib, Bull. Misc. Inform. Kew. 10: 403 (1912); Loesener in Nat.
Pflanzenfam.lSa: 588 (1930). - TYPE; Thailand, Chiang Mai, Doi Sootep, in
evergreen jungle, 660 m., 24 July 1910, Kerr, A.F.G. 1290 (lectotype K! [K000255235],

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Card. Bull. Singapore 67(1) 2015

designated here; isolectot 5 q)es BK n.v., BM! [BM000858181], E! [E00097850], K!
[K000255234], P! [P00450941]).

Zingiber rnenghaiense S.Q.Tong, Acta Phytotax. Sin. 25 (2): 145-146, pi. 1, f. 2.
(1987); S.Q.Tong in FI. Yunnan 8: 533 (1997); T.L.Wu & K.Larsen in FI. China. 24:
326 (2000), synon. nov. - TYPE: China, Yunnan Province, Xishuangbanna Daizu
Zizhizhou, Menghai Xian, Menghai Zhen, under the forest on the roadside, 1 200 m, 2
Jul 1982, Tong, S.Q. &. Li, AM. 32(5(56^(holotypeHITBC! [catalogue number 048849];
isotype KUN! [catalogue number 0833203]).

Zingiber stipitatum S.Q.Tong, Acta Phy to tax. Sin. 25 (2): 146-147, pi. 1, f. 3. (1987);
S.Q.Tong, in FI. Yunnan 8:532(1 997); T.L.Wu & K.Larsen in FI. China. 24: 326 (2000),
synon. nov. - TYPE: China, Yunnan Province, Dehong Daizu Jingpozu Zizhizhou,
Ruili Shi, Mengxiu Xiang, on the way from Mengxiu Cun to Daoba Zhai, 1200 m, 25
Jul 1983, Tong, S.Q. & Liao, C.J. 24536 (holotype HITBC! [catalogue number 48857];
isotypes FllTBC! [catalogue number 49032], KUN! ([catalogue number 0833208],
pro parte, excluding the leafy shoot), KUN! ([catalogue number 0833209], pro parte,
excluding the leafy shoot).

Feremiial rhizomatous herb 1.2-1.6(-2.5) m tall, glaucous throughout (although not
obviously so when old). Rhizomes fleshy, densely branched, 10-20 mm in diameter,
light brown externally, internally purple-pink when young, orange-yellow when
old; root tubers ovate to fusifonn, c. 2 x 1 cm, externally whitish brown, internally
pale grey. Leafy shoots to 15 in a tuft, with up to 44 leaves when flowering, less
than from the base leafless; leaf sheaths longitudinally striate, tubular at base; ligule
l-4(-6) mm long, bilobed, apices obtuse triangular, base green, turning blackish and
coriaceous when old, with obvious raised veins, upper part hyaline, membranaceous,
densely pubescent, glabrescent when old; petiole to 9 mm long, consisting of pulvinus
only, sparsely pubescent; lamina linear to naiTowly-ovate, 15-34 x 1.8-3(^.5)
cm, glaucous and glabrous on both side, base obtuse to attenuate, apex acuminate.
Inflorescences 1-3, radical; peduncles\2-20 cm long, erect, with pink scales on the
base; spikes fusiform or cylindrical, apices acute, 10-15 x 5-6 cm, fertile bracts each
subtending one flower, broadly obovate or broadly spathulate, slightly longer than
the floral tube, green with purple red apices, 3^.5 x 2-A cm, apices rounded, whole
spikes turning scarlet when fruiting; bracteoles naiTOwly ovate, apex acute, c. 30 x 6
mm, semi-translucent, sparsely pubescent externally, glabrous internally. Flowers 5-6
cm long; calyx tubular, membranaceous, c. 1 cm long, unilaterally split to 4 mm, apex
slightly dentate or nearly truncate, sparsely pubescent outside, glabrous inside, semi-
transhicent;y?ora/ tube creamy white, c. 3 cm long; dorsal corolla lobe narrowly ovate,
c. 18x9 mm, cream- white, concave, glabrous, apex mucronate; lateral corolla lobes
narrowly ovate, c. 18 x 7 mm, creamy, glabrous; labellum obovate with conspicuous
attenuate base, c. 26 x 12 mm, creamy, with purple red patch on the base, apex
emarginate or bifid, margin revolute; lateral staminodes narrowly obovate, nan'owly
oblong or narrowly triangular, c. 18 x 4 mm, basal 'A, or even less, connate to labellum.

Zingiber kerrii and the synon3Tny of Z. menghaiense and Z. stipitatum

139

creamy with two purple-red patches on the base, apex rounded to attenuate. Stamen
c. 17 mm \ong; filament reduced to less than 1 mm, anther c. 11 rmii long, comiective
tissue pale yellow, anther thecae 11 mm long, dehiscing throughout entire length,
pollen pale yellow, anther crest c. 6 mm long, pale yellow. Style filiform, white,
glabrous; stigma funnel form, white, ostiole ciliate. Ovary cylindrical, pale yellow,
trilocular, central placentation, with c. 20 ovules in each locule, c, 5 x 3 mm, densely
pubescent; epigynous glands two, pale yellow, c. 4 mm long, 0.3 mm in diameter, apex
blunt. Fruits obovoid to nearly sphaerical or bluntly trigonous capsules, c. 22 x ] g
mm, sparsely villous, greenish-cream with red tinge externally, red internally; seeds
obovoid, c. 5 X 4 mm, dark brown, glossy, with obvious white hilum on the base; aril
white, sac-like, apex with irregular margin, covering most of the seed.

Distribution. Zingiber kerrii is distributed in India, China, Myanmar and Thailand.
(Fig. 5). According to our photographic records, the species is also present in Laos
(Leong-Skomickova, unpublished).

Ecology. In China, Zingiber kerrii occurs in grassy areas on the forest edge, in bamboo
forest and evergreen broad-leaf forest at altitudes of 700-1300 m.

Provisional lUCN conservation assessment. Least Concern (LC). This species has
a large extent of occurrence (around 230,000 km2) and is Icnown from more than
10 localities of which some are protected. We, therefore, provisionally propose this
species to be Least Concern (LC) according to the latest lUCN criteria (2012).

Additional specimens examined. CHINA; Yunnan: Dehong Daizu & Jingpozu Zizhizhou,
Lianghe Xian, Mengyang Zhen, Dangliang Cun, 26 Sep 2012, Bai, L. 12092603 (IBSC);
ibidem, 17 Aug 2013, Bai, L. 13081701 (IBSC); Ruili Shi, Mengliu Xiang, Mangling Cun,
Moli, 21 Sep 2012, Bai, L. 12092103 (IBSC); Xishuangbanna Daizu Zizhizhou, Jinghong Shi,
Jinuo Xiang, Longpa cun, Zhou, S.S. 3212 (HITBC); Menghai Xian, Menghai Zhen, Manduan
Cun, 14 Sep 2012, Bai, L. 12091402 (IBSC); ibidem, 3 Aug 2013, Bai, L. 13080301 (IBSC);
Mengla Xian, on the road from Mengxing Xiang to Mania Zhen, 1 100 m, 21 Oct 2005, Zhou,
S.S. 3069 (HITBC); Mengla Xian, Ymm Xiang, 700 m, 9 Nov 1959, Pei, S.J. 59-11344 (two
sheets in KUN); Zhenyue Xian (Mengla Xian), 1200 m, Nov 1936, Wang, C. W. 80620 (PE).
THAILAND: Chiang Mai: Chom Thong District, Vachi Ratham Waterfall on Doi Intanon,
650-800 m, 15 Sep 1995, Larsen, K. et ai. 46498 (AAU); Doi Sutep, 1250 m, 30 Jul
1968, Sorenseri, T. et al. 3948 (AAU); Doi Sutep, 1120 m, in 1958, Sorensen,!, et al. 4539
(AAU); Inthanon National Park, 850 m, 22 Jul 1988, Phengklai, C. etal. 6757 (BKF); Mae Sa,
Rock Tower Mountain, 18 Sep 1995, Larsen, K. et al. 46636, (AAU); Pong Pho 12 km of Doi
Chieng Dao, 1200 m, 30 Jul 1968, K. Larsen et al. 2874 (E, AAU, BKF); Pliitsanulok: Chat
Trakan District, Phu Soi Dao National Park, 20 Jul 2006, Poulsen, A.D. & Suksathan, P. 2401
(E); Chaiyaphum: Thep Sathit District, 29 Aug 2001, Pooma, R. et al. 2940 (BKF).

Notes. Zingiber kerrii is similar to Z. laoticum Gagnep. in the entire plant being glaucous
externally (particularly prominent in young shoots, becoming inconspicuous when
older), bracts pale gi*een with pink-red margins and rhizomes and pseudostems purple-
red internally. Zingiber laoticum differs from Z. kerrii by the broader label lum which

140

Card. Bull Singapore 67(1) 2015

Fig. 5. Distribution of Zingiber kerrii Craib, based on herbarium material and literature records.
The type localities of the three names are indicated in red. (a) Zingiber kerrii Craib, (b) Z.
menghaiense S.Q.Tong, (c) Z. stipitatum S.Q.Tong.

is richly mottled in dark purple-brown, and longer (e. 15 mm), entire, membranous
ligules.

ACKNOWLEDGEMENTS. We thank Dr Yunfei Deng for critical reading and providing advice
for the paper, the curators of the herbaria CDBI, EMA, GXMG, GXMI, HGAS, HITBC, IBK,
IB SC, KUN, LBG, PE and SYS for their help with the specimens for our studies, the curators
of AAU, BK, BKF, BM, C, E, G, HAST, K, KFRI, L, P, SING, TAI, TAIF, US, W, WU for
the high-resolution images of specimens. The first author is grateful to the Singapore Botanic
Gardens for granting research fellowships to support the study at SING. The research of the first
and third authors was supported by the National Natural Science Foundation of China (Grant
no. 31170185), the research of the second author is supported by the National Parks Board
Singapore and the Grant Academy of the Czech Republic (GACR grant no. P506/14/13541S).

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Gardens’ Bulletin Singapore 67(1): 143-149. 2015
doi: 10.3850/S2382581215000150

143

Pennilabium labanyaeanum (Orchidaceae), a new species

from Meghalaya, Northeast India

C. Deori, N. Odyuo & A. A. Mao

Botanical Survey of India, Eastern Regional Centre,
Sliillong-793003, Meghalaya, India
drchayadeoribsi@gmail.com
nodyuo@yahoo.com
aamao2008@gmail.com

ABSTRACT. Anew orchid species, Pennilabium labanyaeanum C.Deori, N.Od5aio & A. A.Mao,
is described and illustrated from Meghalaya, India. A key to the species of Pennilabium in India
is provided.

Keywords. India, Meghalaya, new species, Orchidaceae, Pennilabium

Introduction

The genus Pennilabium JJ.Sm. (Orchidaceae) (penni- = feathered, -labium = lip) is a
genus of 1 6 species of epiphytes in the tribe Vandeae, subtribe Aeridinae. It is distributed
from northeastern India and southern China to the Philippines and Sulawesi, with
centres of diversity in Peninsular Malaysia and Borneo (Seidenfaden, 1988; O ’Byrne,
2000; O’Byme et ah, 2009; Schuiteman, 2013; Govaerts, 2014; Pridgeon et al., 2014).
The plants are small twig or branch epiphytes, the flowers are ephemeral and open in
succession, and the lip is spurred with small or large and often fimbriate or marginally
toothed side lobes, often with a fleshy mid-lobe. The column lacks a foot and contains
two pollinia on a long strap-like stipe; the rostellum is nonnally beaked (Pridgeon et
ah, 2014).

In India two Pennilabium species are known, both of which are confined to
the Northeast: namely, P. prohoscidcum A.S.Rao & J.Joseph from Meghalaya and
P. struthio Carr from Arunachal Pradesh. During a botanical tour to Laitkyrhong,
5 km from Smith, East Kliasi Hills District, Meghalaya, in July 2014, the first two
authors discovered a small Pennilabium with pure white flowers. After critical study
of specimens and the literature (Holttum, 1957; Rao & Joseph, 1969; Garay, 1972;
Comber, 1990; Rao, 1996), and comparison to the protologue and spirit material
of the type specimen of the morphologically most similar taxon, Pennilabium naja
P.O’Byme (O’Byme, 2000), this plant was found to be morphologically different from
all other species. Our plant from Meghalaya is therefore described as the new species
Pennilabium labanyaeanum C.Deori, N. Odyuo & A. A.Mao. Schuiteman (in Pridgeon
et al., 2014: 230) points out that ‘the generic delimitation of the genera closely allied
to Pteroceras, including Pennilabium, needs further study, and Pennilabium is only
tentatively accepted here’. Kocyan & Chase (in Pridgeon et al., 2014: 230) pointed

144

Card. Bull. Singapore 67(1) 2015

that only two Pennilabium species were sampled in their molecular study, and said ‘it
would appear that some changes to the taxonomy of this clade will be required once
greater sampling of the species has been accomplished.’

Pennilabium labanyaeanum C.Deori, N.Od 5 uio & A.A.Mao, sp.nov.

Pennilabium labanyaeanum is similar to P. naja in having reduced sidelobes but
differs in the flower being only half open; the sepals and petals pure white; the sepals
obtuse, keeled dorsally and 5-veined; the lip base forming a narrow hollow channel
with minute hairs; the lip epichile imlobed with erose margins, without a prominent
raised callus but with an area of yellow powdery substance widening towards the
lip apex; and the short white spur (see Table 1, Fig. 2). - TYPE; India, Meghalaya,
Laitkyrhong, 5 km from Smith, East Khasi Hills, 1753 m, 25°26'53.2"N 91°52'48.6"E,
23 July 2014, Deori, C. & Odyuo, N. 131601 (holotype CAL [as 131 601 A]; isotypes
ASSAM [as 131601 B-E]), (Fig.l, 2)

Small monopodial epiphytic herbs, 3-5 cm tall, glabrous. Roots 5-30 cm long, 2-4
mm in diameter, several from base of stem below leaves, white with green tips. Stems
1-2.5 X 0. 3-0.5 cm, very short, compressed with a few leaves close together. Leaves
1.5-6 X 0.5-1. 5 cm, distichous, elliptic or broadly lanceolate, more or less fleshy,
often twisted at the base, leathery, green, basally sheathing, unequally bilobed at apex,
apices subacute to obtuse. Inflorescence 2.5-6 cm long, unbranched, lateral, emerging
through sheath opposite leaf, erect or parallel to the leaves, producing 2-3 flowers
in succession; peduncle 2-3.5 cm long, slender, cylindrical, base covered with two
sheaths; sheathing sterile bracts below rachis two, c. 2 x 4.5 mm, somewhat rectangular
in shape, lemon green; rachis 5-10 mm long, complanate-cylindrical, ridged with one
to three distichously arranged flowers opening in succession; floral bract 2-2.5 x c.
1.5 mm, persistent, ovate-triangular, obtuse, lemon green. Flowers 12-14 mm long
from the tip of the dorsal sepal to the tip of the spur, 8-10 rmii broad, ephemeral,
half open, formed in succession, sepals and petals pure white with yellow markings
on the epichile of the lip. Pedicel with ovary 1-2.6 cm long, lemon green, 3 -ridged.
Sepals oblong with narrow bases, obtuse, 5-veined, on the dorsal side with keels which
apically project as small apicules; dorsal sepal 7-10 x 3-4 mm; lateral sepals 8-9.5 x
3.5^ mm. Petals 7.5-9 x 2.5-3 mm (base nan-ow), oblong with narrow base, obtuse
with 3 main veins, lateral veins branched. Lip spuiTed, 1 1.5-12.5 mm long (including
the spur), 4-5 rmu wide at the centre and 3 mm at the apex, unlobed, suborbicular or
ovate, lateral margins of lip erose; lip base with a naiTow hollow chamiel with minute
hairs within; epichile fleshy, solid, thick (1 mm in diameter), with an area of yellow
powdery substance widening towards the lip apex. Spur 5-7 mm long from junction
of lip base and the column, white, slender, tip rounded, 1.5 mm wide, aseptate and
ecallose; longitudmal section of the spur shows sparse minute hair-like structure or
raised edges at mouth towards the centre. Column 2-2.5 x c. 2 mm, without a foot,
rounded, without stelidia; stigmatic cavity very broadly ovate; rostellum not seen.
Anther 1-1.5 x 1.5-2 mm, operculum c. 0.2 x 0.5 mm, truncate, entire. Pollinia two.

Pennilabium labanyaeanum, a new orchid from India

145

Fig. 1. Pennilabium labanyaeanum C.Deori, N.Odyiio & A. A. Mao. A-C. Habit. D.
Inflorescence. E. Flower bud. F. Flower (side view). G. Peduncle sheath. H. Floral bract. I-J.
Flower (front & side views). K. Floral perigon with spurred lip, L. Column with pedicel and
ovary. M. Dorsal sepal (dorsal view). N. Lateral sepal (dorsal view). O. Lip (front view). P. Lip
front view (highlighting the narrow hollow channel with minute hairs at the base). Q. Spurred
lip (side view) with column, pedicel and ovaiy. R. Longitudinal section of spur. S-T. Anther
(front views). U. Pollinia with stipe. V. Stipe. W. Fruit. (Drawn by: C. Deori)

146

Card. Bull Singapore 67(1) 2015

Pennilabium labanyaeanum, a new orchid from India

147

c. 1 mm long (including stipe), 0.5-0. 7 mm in diameter, globose, entire; stipe 0.7-1
mm long, widening near the attaclunent of the pollinia, with a small disc. Fruits erect,
slender, 3-5.2 cm long which is more than twice the flower length, 3-ridged.

Distribution and habitat. Pennilabium labanyaeanum was found growing epiphytically
on moss-covered branches of trees. The species was flourishing in a steep subtropical
hill forest along a stream side at 1753 m altitude near Laitkyrhong, 5 km from Smith,
East Kliasi Hills District, Meghalaya, India. It was associated with other orchid species
in the genera Dendrobium Sw. and Eria Lindl. s.l., amongst others.

Phenology. Flowering and fruiting: July-August.

Etymology. The speciflc epithet is given in honour of the first author’s mother (L.
Labanya Deori, 1945-1993).

Provisional lUCN consen’ation assessment. Data Deficient (DD). Pennilabium
labanyaeanum is so far known only from a single locality in the East Khasi Hills District
in Meghalaya. According to O’Byme (2009), all species in the genus Pennilabium are
uncommon or rare and occur in small colonies that can be widely separated from each
other. If this is also the case for Pennilabium labanyaeanum, the new species might
be more widespread in Meghalaya than is currently known and further exploration is
necessaiy before a threat status can be given.

Notes. Pennilabium labanyaeanum is distinct in the genus due to its unlobed
suborbicular lip. The new species is similar to Pennilabium naja from Sulawesi
but differs as discussed above (see also Table 1, Fig. 2). As already pointed out by
Garay (1972) there are two types of lip stmcture in Pennilabium: 1 . Lateral lobes well
developed and midlobe reduced to a small fleshy lobe; 2. Lateral lobes much reduced,
possibly represented only by small auricle-like lobes, and midlobe very prominent.

Fig. 2. A-S. Pennilabium labanyaeanum C.Deori, N.Odyuo & A.A.Mao. A-A2. View of the
type locality. B-B2. Pennilabium labanyaeanum in the wild. C-C2. Habit. D. Inflorescence.
E. Floral bract. F. Peduncle sheath. G & Gl. Flower, front & side views. H. Fruit. 1. Dorsal
sepal, dorsal view. J. Lateral sepal, dorsal view. K. Flower bud, side view. L. Lip, front view.
LI. Lip, front view (highlighting the narrow hollow channel with minute hairs at the base).
M. Floral perigon with spurred lip. N. Spurred lip(side view) with column, pedicel and ovary.
O. Column, close view. Ol. Column with pedicel & ovary. P & PI. Anther, front views. Q.
Longitudinal section of spur. R. Pollinia with stipe. S. Stipe. Pennilabium naja P.O’ Byrne
(images taken from the spirit sample of the holotype O 'Byrne SUL107 at SING) T. Flower, side
view. U. Floral perigon with spurred lip. V. Lip, front view. W. Anther, front view. X. Pollinia
with stipe. (Photos: C. Deori & N. Odyuo)

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Card. Bull. Singapore 67(1) 2015

Table 1. Comparison of Pennilabium naja and P. labanyaeamim.

Pennilabium naja

Pennilabium labanyaeanum

The flowers are fully open with yellow
sepals and petals, about 12 mm wide.

The flowers are not fully open with
pure white sepals and petals, 8-10 mm
wide.

Sepals spreading with recurved tips,
oblong obtuse, 3 -veined, not keeled.

Sepals not spreading, broadly oblong
obtuse, 5 -veined, keeled dorsally
at the apex.

Lip spurred, fleshy, tongue shaped,
white with lateral wings at the base
of the spur aperture; side lobes not
distinct; midlobe ovate in outline,
recurved in centre, epichile obtuse,
lamina concave with rolled under
margins, fleshy, iTigulose, raised into a
broad low ridge along median line from
base of callus to just before lip apex.

Lip spurred, fleshy, sub-orbicular, white
with no lateral wings; sidelobe absent;
midlobe not recurved at the centre,
epichile unlobed with erose margins,
without a prominent raised callus
but with an area of yellow powdery
substance widening towards the lip
apex.

Spur yellow, 9-10 mm long, slender.

Spur white, 5-7 mm long.

fleshy and solid. Most of the species in the genus fall into the first category. Two of
the three species found in India, Pennilabium proboscidcum and P. struthio, have well
developed side lobes, but the lip of the new species R labanyaeanum lacks side lobes
altogether, thereby separating it from all other species in the genus.

A key to the species of Pennilabium in India

la. Lip sidelobes present, large and fimbriate 2

lb. Lip sidelobes absent P. labanyaeanum

2a. Sepals and petals purple dotted, creamy white; lip sidelobes at least twice as

long as broad at the truncate apex, which has erose-fimbriate edges

P. proboscidcum

2b. Sepals and petals yellowish with dark red; lip sidelobes not twice as long as

broad near the truncate apex, which has a rim of dense, branching lacinia

P. struthio

Pennilabium labanyaeanum, a new orchid from India

149

ACKNOWLEDGEMENTS. The authors are thankful to Dr P. Singh, Director, Botanical Survey
of India for encouragement and facilities. The authors are thankful to Dr Andre Schuiteman,
Senior Researcher, Herbarium, Royal Botanic Gardens, Kew, for his comments in support
of the novelty of the species described here. We are also grateful to Guat Bee from OSSEA
(Orchid Society of Southeast Asia) for sending the protologue of Pennilabium naja and to
Bazilah Ibrahim of the Herbarium at Singapore Botanic Gardens, for sending spirit material of
the holotype of Pennilabium naja for study. We are also thankful to Rudan Kliakhongor and
Iboklang Kharhongor for assisting us during the field trip.

References

Comber, J.B. (1990). Orchids of Java. UK: Royal Botanic Gardens, Kew.

Garay, L. (1972). On the systematics of the monopodial orchids I. Bot. Mus. Leaf. 23(2):
149-212.

Govaerts, R., Bemet, R, Kratochvil, K., Gerlach, G., Can-, G., Alrich, R, Pridgeon, A.M., Pfahl,
J., Campacci, M.A., Holland Baptista, D., Tigges, H., Shaw, J., Cribb, R, George, A.,
Kreuz, K. & Wood, J.J, (2014). World Checklist of Orchidaceae. Royal Botanic Gardens
Kew. http://apps.kew.org/wcsp/home.do (25 Jul. 2014).

Holttum, R.E. (1957). Flora of Malaya (Orchidaceae), vol.l. Singapore: Government Printing
Office.

O’Byme, RO. (2000). Three new orchid species from Sulawesi. Malayan Orchid Rev. 34: 64.
O’Byme, RO., Phoon, S.N. & Ong. RT. (2009). Pennilabium armariv. A new species from
Peninsular Malaysia. Malesian Orchid J. 3: 113-116.

Pridgeon, A.M., Cribb, P.J., Chase, M.W. & Rasmussen, F.N. (2014). Genera Orchidacearum,
vol. 6 (Epidendroideae, part 4). UK: Oxford University Press.

Rao, A.S. & Joseph. J, (1969). Pennilabium proboscidcum A.S. Rao & Joseph — Anew orchid
species from K. & J. Hills, Assam with incidental first record of the genus from India.
Bull. Bot. Surv India 10: 231.

Rao, A.N. (1996). Five new records of orchids from Arunachal Pradesh, India. J. Econ. Taxon.
Bot. 20(3): 708.

Schuiteman, A. (2013). Avery odd thing: Pennilabium hewittii (Orchidaceae). KeM> Bull. 68:
679-681.

Seidenfaden, G. (1988). Orchid genera in Thailand XIV. Fifty-nine Vandoid Genera. Opera
Bot. 95: 330-332.

Gardens’ Bulletin Singapore 67(1): 151-157. 2015
doi: 10.3850/S2382581215000162

151

Tripogon bimucronatus (Poaceae: Chloridoideae:
Tripogoninae), a new species from India

K. Thoiba^, A.K. Pradeep^ & C.N. SuniP

^Department of Botany, University of Calicut,
Kerala, 673635, India
akpradeep 1 @rediflFmail.com

^Department of Botany, S.N.M. College, Maliankara,
Kerala, 683516, India

ABSTRACT. Tripogon bimucronatus Tlioiba & Sunil, a new species of grass from the
Nelliyampathy Hills, Palakkad District of Kerala, South India is described and illustrated. It
is moiphologically similar to Tripogon vellarianus Pradeep but differs by having adaxially
scabrid, linear-acuminate leaves; ligules a fringe of hairs to 2.5 nun long; spikelets with 6-8
closely packed florets; lower glume asynuuetrical; upper glume awned; lemma awn scabrid;
palea elliptic-lanceolate, apex bimucronate and notched at the centre.

Keywords. Tripogon, Poaceae, new species, Kerala, South India

Introduction

The genus Tripogon Roem. & Schult. belongs to subtribe Tripogoninae Stapf, tribe
Cynodonteae Dumort, subfamily Chloridoideae Kunth ex Beilschm. in the family
Poaceae Barnhart (Peterson et al., 20 1 0, 20 14; Soreng et al., 20 1 2). The genus comprises
44 species distributed in Africa, America, Australia, Temperate and Tropical Asia
(Clayton et ah, 2006 onwards). It is known to have 22 species in India, which includes
six recently described species (Murugesan & Balasubramaniam, 2008; Newmaster et
al., 2008; Kabeer et al., 2009; Chorghe et al., 2013; Thoiba & Pradeep, 2014).

During an exploration along the Western Ghats of South India, an interesting
specimen of Tripogon was collected from the hill slopes of the Nelliyampathy, Palakkad
District of Kerala, South India. Critical study revealed it to be quite distinct from the
remaining species of the genus and hence is described here as a new species, bringing
the total number of Tripogon species for India to 23. It is closely allied to Tripogon
vellarianus Pradeep (Pradeep & Sunil, 1999) but can easily be distinguished (Table 1).

Tripogon bimucronatus Thoiba & Sunil, sp. nov.

It is very similar to Tripogon vellarianus Pradeep but differs in having adaxially scabrid,
linear-acuminate, ribbed leaves, ligules being hairy at the ends of leaf sheaths, with
the hairs reaching up to 2.5 mm; spikelets with 6-8 closely packed flowers; the lower
glume being asymmetrical, and the apex of upper glume awned; lermna 1 -awned,
scabrid, straight or geniculate; palea elliptic-lanceolate, apex bimucronate and notched

152

Card. Bull. Singapore 67(1) 2015

at the centre. - TYPE: India, Kerala, Palakkad District, Nelliyampathy hill top, 1200
m elevation, 16 November 2010, Sunil, C.N. 4477 (holotype SING; isotypes BRIT,
CALI,K). (Fig, 1,2)

Tufted perennial herb. Culms 30-60 cm tall, erect; nodes glabrous. Leaf blades
30-60 cm long, 0.4-0.6 cm wide, linear, hispid towards base, apex acute-acuminate,
scabrid adaxially, margins involute. Sheaths closely involute to the culm, almost rigid,
sparsely hairy; ligtiles a fringe of hairs 2-2.5 mm long. Racemes 30^0 cm long,
spikelets arranged along both sides, 45-65 spikelets per raceme; peduncle glabrous,
terete, 10-15 cm long; rachis stout, minutely scabenilose, triangular. Spikelets linear-
lanceolate, 1 .3-1.8 cm long, 2-2.5 mm wide, 6-8-flowered; callus bearded. Rachilla not
persistent, 1-2 mm long, slightly scabrid. Lower glume 3-3.5 x 0.5-1 mm, lanceolate,
asymmetrical, notched on one-side below the middle, chartaceous, 1 -nerved, apex
acuminate. Upper glume 5-6 x 0.7-1 mm, elliptic-lanceolate, prominentely 3-nerved,
glabrous, apex with awns 0.5-1 mm long. Lemma 5-5.5 x 1 .5-2 mm (excluding awn),
3-nerved, l-awned, the awn arising between lateral lobes, median awn 5-5.5 mm long,
scabrid, straight or geniculate, lateral awns absent. Palea 4-5 x 1-2 rmu, hyaline,
narrowly elliptic, keeled and winged, keels minutely puberulous, apex bimucronate,
notched at the centre. Lodicules 2, c. 0.25 mm, quadrate, apex coarsely 3-toothed.
Stamens 3, anthers 1-1.5 mm long, oblong, filaments 0.5-0.75 mm long, slender,
glabrous. Ovary 0.25-0.5 mm long, obovate; style 2, slender, hyaline, 1 mm long;
stigma feathery, 1-1.5 mm long. Caryopsis 1. 1-1.3 x 0.4-0. 5 mm, naiTowly oblong-
lanceolate, obtusely trigonous, pale brown, obtuse at apex, longitudinally ribbed
abaxially.

Distrihution & Ecology^ Tripogon bimucronatus usually grows in the granitic grassy
slopes of Nelliyampathy Hills at an elevation of 1000-1200 m. It is found growing in
association with Arthraxon castratus (Griff.) V. Naray. ex Bor, Garnotia elata (Am.
ex Miq.) Janowsky., Cyanotis papilionacea (Bunn.f.) Schult. & Schult.f, Swertia
angustifolia Buch.-Ham. ex D. Don., Isachne bourneorum C.E.C.Fisch., Tripogon
wightii Hook.f., Tephrosia pulchella Hook.f, Cymbopogon commutatus (Steud.) Stapf,
Osbeckia spp., Parasopubia delphiniifolia (L.) H.-P.Hofm. & Eb, Fisch., Chrysopogon
nodulibarbis (Hochst. ex Steud.) Henrard, Sopubia trifida Buch.-Ham. ex D.Don,
Eulalia trispicata (Schult.) Henrard, and various moss species. It also occurs along
Meenuliyan Para Hills in Idukki District, Kerala at an elevation of 800 m.

Phenology^. It flowers from early September to October. Fmit set is low with the seeds
ripening by late October-November.

Etymology. The epithet ‘bimucronatus’ refers to the bimucronate palea apex.

Additional specimen examined. INDIA: Kerala: Idukki Dt., Meenuliyan Para, Vannappuram,
800 m elevation, 28 Nov 2014, Sunil 4910 (SNM College Herbarium, Kerala).

Tripogon bimucronatus, a new grass from India

153

Table 1. Comparison between Tripogon vellarianus Pradeep and Tripogon bimucronatus
Thoiba & Sunil.

Tripogon vellarianus

Tripogon bimucronatus

Leaf blades 8-15 mm wide, glabrous
on both surfaces

Leaf blades 5-8 mm wide, ribbed,
slightly scabrid adaxially

Leaf sheaths closely clasping, very
rigid; ligules membranous

Leaf sheath closely involute, almost
rigid; ligules ciliate, 2-2.5 mm long

Spike] ets distantly arranged,
8-10-flowered

Spikelets closely packed,

6-8-flowered

Lower glumes 4-5 x l .25 mm,
symmetrical, unlobed, acuminate at
the apex

Lower glumes 3-5 x 1mm,
asymmetrical, slightly 1-lobed on
one-side, acute-acuminate at the apex

Upper glumes 5-7.5 x 1.5 mm, acute
at apex, unawned

Upper glumes 5-6 x 1mm, a small
awn between the sinus at apex, awns

0.5-1 mm long

Lemmas 5-10 x 2 mm (excluding
awn), lateral awns absent or up to

5mm long, median awn 3 mm long,
glabrous, straight

Lemma 5-5.5 x 1 -2mm (excluding
awn),l-awned in between the lobes,
awn 5-5.5 mm long, scabrid, straight
or geniculate

Paleas c. 4 x 2 mm, elliptic, acute at
apex

Paleas 4-5 x 1-1.2 mm, elliptic-
lanceolate, bimucronate at apex with a
notch at the centre

Key to the species of Tripogon in India
(with lemma entire or 2-cleft at apex)

la. Lowest lemmas empty, dissimilar to other lemmas T. wardii

lb. Lowest lemmas bisexual, similar to all other lemmas 2

2a. Culms thickened below by the persistent leaf sheaths; leaf blades equitant, rigid,

apex pungent T. pungens

2b. Culms not thickened below by the persistent leaf sheaths; leaf blades not equitant,
not rigid or pungent 3

Tripogon bimucronatus, a new grass from India

155

3a. Central awns of the lemmas flexuous, capillary, several times as long as lemmas

T. capillatus

3b. Central awns of the lemmas straight or curved, not more than twice as long as the

lemmas 4

4a. Rachilla intemodes 1 mm long; apices of lemmas 1-awned 5

4b. Rachilla intemodes 2 mm long; apices of lemmas 3-awned T. copei

5a. Culms 15-25 cm high; leaf blades 5-20 cm long 6

5b. Culms 30-90 cm high; leaf blades 30-60 cm long 7

6a. Upper glumes 5.5-7 mm long; awns of the lemmas 6-8 mm long, straight or

curved T. wightii

6b. Upper glumes 8-9.5 mm long; awns of the lemmas 10.5-12 mm long, always
straight T. velliangiriensis

1 a. Leaf blades and culms glaucous, involute, filifomi; ligules very short and ciliate ....

T. jacquemontii

7b. Leaf blades and culms green, flat, sometimes rolled; ligules obsolete 8

8a. Leaf blades 3-6 mm wide; racemes 7-25 cm long; median awns shorter than

lemmas T. lishoae

8b. Leafblades 8-15 mm wide; racemes 30^0 cm long; median awn equal to or longer
than lemmas 9

9a. Ligules membranous, a fringe of hairs at the ends of leaf sheath; paleas subulate at
apex T. malabaricus

9b. Ligules ciliate, membranous or absent; paleas not subulate at apex 10

10a. Upper glumes 3-4 mm long; lemmas 2. 5-3. 5 m i n long 11

10b. Upper glumes 4-7.5 mm long; lemmas 4-10 mm long 12

lla. Ligules glabrous; lemma base glabrous, median awns as long as or shorter than the

lemmas; inflorescence 20-45 cm long; spikelets with 5-8 florets

T. sivarajanii

llb. Ligules ciliate at apex; lemma base bearded; median awns 1.5 times longer than
lemma; inflorescence 15-20 cm long; spikelets with 9-10 florets ... T. tirumalae

12a. Ligules a fringe of hairs; upper glumes awned at sinus; apex of paleas bimucronate

T. bimucronatus

12b. Ligules glabrous; upper glumes acuminate at apex; apex of paleas acute

T. vellarianus

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Card. Bull. Singapore 67(1) 2015

Fig. 2. Distribution map of Tripogon vellarianus Pradeep (■) and Tripogon bimucronatus
Thoiba & Sunil (•) in Kerala

ACKNOWLEDGEMENTS. The authors are thankful to Dr K.H. Amitha Bachan, Western
Ghats Hombill Foundation (WGHF), Thrissur, Kerala for help during collection trips to the
Nelliyampathy hills, Palakkad District of Kerala, and to the anonymous reviewers for useful
suggestions that improved the text. The last author is thankful to the University Grants
Commission, New Delhi for financial assistance.

References

Chorghe, A., Rasingam, L., Prasanna, P.V. & Rao, M.S. (2013). Tripogon tirumalae (Poaceae),
a new species from the Seshachalam Hills of Andhra Pradesh, India. Phytotaxa 131:
17-22.

Clayton, W.D., Vorontsova, M.S., Harman, K.T. & Williamson, H. (2006 onwards). GrassBase-
the online world grass flora, http://www.kew.org/data/grass.db.html. (accessed on 10
Aug. 2013).

Tripogon bimucronatus, a new grass from India

157

Kabeer, K.A.A., Nair, V.J. & Muithy, G.V.S. (2009). Tripogon borii - a grass species new to
science from India. Bull. Bot. Siirw India 50: 1 15-118.

Murugesan, M. & Balasubramaniam, V. (2008). Tripogon velliangiriensis (Poaceae) - a new
species from Tamil Nadu, India. Indian J, Forest. 31: 109-111.

Newmaster, S.G., Balasubramaniam, V., Murugesan, M. & Ragupathy, S. (2008). Tripogon
cope (Poaceae: Chloridoideae), a new species supported by morphometric analysis and
a synopsis of Tripogon in India. Syst. Bot. 33: 695-701.

Peterson, P.M., Romaschenko, K., & Jhonson, G. (2010). A classification of the Chloridoideae
(Poaceae) based on multi-gene phylogenetic trees. Molec. Phylogenet. Evol. 55: 580-
598.

Peterson, P.M., Romaschenko, K. & Herrera Arrieta, Y. (2014). A molecular phylogeny and
classification of the Cteniinae, Farragininae, Gouiniinae, Gymnopogoninae, Perotidinae,
and Trichoneurinae (Poaceae: Chloridoideae: Cynodonteae). Taxon 63:275-286. http://
dx.doi.org/1 0. 1 2705/632.35

Pradeep, A.K. & Sunil, C.N. (1999). Two new species of Tripogon (Poaceae) from India. Sida
18: 809-814.

Soreng, R.J., Davidse, G., Peterson, P.M., Zuloaga, F.O., Judziewicz, E.J., Filgueiras, T.S.,
Mon'one, O., & Romaschenko, K. (2012). A world-wide phylogenetic classification of
Poaceae (Graminae). http://www.tropicos.org/projectwebportal.aspx?pagename=Classi
ficationNWG&projectid=10 (accessed on 10 Aug. 2013).

Thoiba, K. & Pradeep, A.K. (2014). Tripogon malabarica (Poaceae: Chloridoideae:
Tripogoninae), A new species from India, Kerala. J. Bot. Res. Inst. Texas 8: 525-529.

Gardens’ Bulletin Singapore 67(1): 159-229. 2015
doi: 10.3850/S2382581215000174

159

A revision of Epithema (Gesneriaceae)

K. Bransgrove^’^ & DJ. Middleton^

'Royal Botanic Garden Edinburgh, 20A Inverleith Row,
Edinburgh, EH3 SLR, Scotland, U.K.

^Herbarium, Singapore Botanic Gardens, National Parks Board,

1 Cluny Road, Singapore 259569
david_niiddleton@nparks.gov.sg
^Current address: Australian Tropical Herbarium,

James Cook University, Cairns Campus, Building E2,

RO. Box 6811, Cairns, Queensland 4870, Australia

ABSTRACT. The genus Epithema Blume (Gesneriaceae) is revised. A key is given, all
species are described and several are illustrated. Twenty species are recognised and two
new combinations are made, Epithema philippinum (Hilliard & B.L.Burtt) Bransgrove and
Epithema pusillum (C.B. Clarke) Bransgrove. Provisional lUCN conservation assessments are
given for all species.

Keywords. Africa, Epithema, Gesneriaceae, Southeast Asia, taxonomic revision

Introduction

Epithema Blume is a genus of lithophytic herbs in central tropical Africa (Cameroon,
Central African Republic, Democratic Republic of Congo, Equatorial Guinea, Guinea,
Ivory Coast, Liberia, SieiTa Leone and Uganda), South Asia (Nepal, India, Sri Lanka),
Southeast Asia (southern China, Myanmar, Thailand, Vietnam, Cambodia, Laos,
Taiwan), Malesia (Philippines, Malaysia, Indonesia) and the Solomon Islands.

The genus was first described by Blume (1826) with a single species, Epithema
saxatile Blume. Aikinia R.Br. (in Wallich, 1832) was published shortly thereafter for a
different species {Aildnia brunonis) but it was soon synonymised under Epithema by
Decaisne (1834). Bentham (1835) and Spanoghe (1841) added further combinations or
new taxa to Epithema but Don (1838) continued the use of Aikinia, placing Epithema
in synonymy and moving Epithema carnosiim to Aikinia. Later authors, however,
correctly published treatments under the name Epithema. Major publications on the
genus mclude two complete treatments of Epithema (Candolle, 1845; Clarke, 1883).
Candolle (1845) included five species in his account and Clarke (1883) six species, of
which five were from Asia and one from Africa. The account by Clarke (1883) also
included eight new varieties and one new varietal combination. After 1883 there were
occasional additional regional accounts and the descriptions of new taxa (Henriques,
1892; Chevalier, 1912; Merrill, 1916; Ohwi, 1943; Burtt, 1958; Kiew, 1985; Ying,
1992; Li & Kao, 1998; Wang et al., 1998) and a set of brief but valid publications
of nine new taxa and combinations by Hilliard & Burtt (1997) in anticipation of a
revision of the genus which, sadly, never materialised.

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Card. Bull. Singapore 67(1) 2015

The classification of the Gesneriaceae into subfamilies, tribes etc. has been
rather unsettled (Weber et ah, 2013) with the result that Epithema has been placed in a
number of different subfamilies, tribes and series. Burtt & Wiehler (1995) and Weber
et al. (2013) both placed Epithema in the tribe Epithemateae (Meisn.) C.B.Clarke of
the subfamily Didymocarpoideae (called Cyrtandroideae by Burtt & Wiehler (1995)).
Weber et al. (2013) further placed it in its own subtribe, Epitheminae. Mayer et al.
(2003), in a molecular phylogenetic study of the group, found that the Epithematoid
Gesneriaceae are sister to the Didymocarpoid Gesneriaceae and that Epithema is
monophyletic and sister to a clade with a single species each of Loxonia Jack and
Stauranthera Benth. The study only included four species of Epithema {E. tenue, E.
saxatile, E, membranaceum and E. taiwanense [= E. ceylanicum]) and, therefore,
provides little information about the interrelatedness of the species within the genus.

In flower, Epithema is not easily confused with other genera although some
specimens have been previously mistaken for MonophyUaea R.Br. due to a superficial
resemblance to some species. The two genera can be separated by inflorescence
arrangement, sepal shape and aestivation, placentation, the number of locules in the
ovary, seed shape and hair type.

The species of Epithema are not readily distinguished from each other. Clarke
(1883) suggested that the then known five Asian species could possibly be recognised
as only one. Many are similar and there are few good characters to separate the species
as both vegetative and reproductive characters can be variable. In particular, the species
from mainland Asia are very similar and difficult to separate.

The gender of Epithema is neuter. Clarke (1883), however, described several
new varieties with feminine endings. These have been corrected to neuter in the
revision below.

Morphological Characters

Epithema is primarily lithophytic. In eastern and southeastern Asia it is found on
limestone and, to a lesser extent, on granite and sandstone. It appears to be restricted
to limestone in Borneo. In Africa, India and non-peninsular Myamnar it is found on
volcanic rocks and soils. It requires shaded conditions with high humidity and is
frequently found near or in cave entrances, on rocks on the forest floor, or on rocks
beside, or in, shallow streams and rivers. Information from specimen labels indicates
Epithema species will occasionally grow on fallen trees and is often collected from
road sides and road side embankments.

All species of Epithema are small herbs. Epithema, like other didymocarpoid
Gesneriaceae, is anisocotyledonous with the cotyledons withering early. The lowermost
true leaf, however, is solitary and looks rather like the macrocotyledon found in other
genera such as MonophyUaea (Weber, 2004).

The majority of species are caulescent but a few species have a very short
stem with leaves arising more or less at ground level, therefore appearing acaulescent
(Epithema dolichopodum, E. Jongipetiolatum, E, philippinum, E, rennellense, E.
strigosum and some specimens of E. benthamii, E. ceylanicum and E. tenue). One

Revision of Epithema

161

caulescent species, however, often exhibits a more sprawling rather than an upright
habit {E. sarawakense) and one is unifoliate in most of its distribution (E. horsfieldii).

The hairs on all plant parts are simple, unbranched, straight or hooked, have 3-5
cells and are from 0.02 mm to (rarely) 1.5 mm long. Hooked hairs are usually shorter
than straight hairs. The type of hairs (pubescent, strigose, setose, hispid or villous),
density (glabrous to dense), colour (hyaline to brown) and length are quite variable.
The combination of these characters is useM in some species but in others it is often
too variable to be of any taxonomic use. It is, therefore, included in the descriptions
only where it is taxonomically useful. Where ‘minute hooked pubescence’ is included
in descriptions it denotes hooked hairs of a maximum of 0.1 mm long.

The leaves of Epithema are thinly to thickly membranous and are usually
narrowly to broadly ovate, more rarely oblong to linear. The apex of the leaves is
rounded to acute, the margin entire to serrate, dentate or crenate (or variations thereof)
and the bases cordate to truncate to occasionally attenuate. The upper surface of
the leaves is green to dark or black-green and the lower surface is usually lighter in
colour than the upper surface and may dry whitish green to grey-green. Specimens of
Epithema saxatile and E. ceylanicum, and possibly others, show some green and silver
variegation on the upper surface of the leaves. The variegation has only been seen in a
small number of specimens from Thailand and Peninsular Malaysia. The indumentum
of the upper surface is variable in composition and the hairs are up to 1.5 mm long. The
indumentum of the lower surface is similar to that of the upper surface but the hairs
are usually sparser, shorter and finer. The indumentum of the leaves is occasionally
taxonomically useful and is only included in the descriptions in those cases.

The leaf venation is cladodromous with the secondary veins mostly strongly
ascending. There are often weaker intersecondaries and the tertiary venation is
mostly fairly obscure but, when visible, laxly alternate percurrent. The number and
arrangement of secondary veins is variable, between 4 and 1 5 and alternate to opposite
(or both witliin one leaf), and not taxonomically important.

Leaf arrangement is variable. Caulescent species can have opposite or solitary
leaves that in turn may be petiolate or sessile. A combination of all leaf an*angements
may be found on one plant or within a population. Most acaulescent species have
unpaired leaves. When present, solitary upper leaves are nearly always petiolate and
opposite one or more peduncles.

Weber (2004) described the infiorescence of Epithema as “reduced to a single
bract/inflorescence unif ’ which is derived from a “more elaborate thyrse, with several
bract/inflorescence units in alternate position”. The appearance is of a small, multi-
flowered, scorpioid cyme. The flowers are an*anged into rows, two to thi*ee across.
The indumentum of the peduncle is glabrous to hispid with hairs to 1 .25 mm long.
The flowers are collectively subtended by one bract which is green and frequently
cucullate, partially or entirely enclosing the inflorescence, but can be very small. The
uppermost margm of the bract is usually dentate but can be entire to dentate. If the
margin of the bract is lobed, the margin of the lobes may be entire, dentate or crenate.
The indumentum of the lower surface of the bract is of a short, hooked pubescence
and/or strigose to setose, straight and/or hooked hairs to 1 mm long. The indumentum

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Card. Bull. Singapore 67(1) 2015

of the upper surface is usually of one hair type only, towards the margin and to 0.6
mm long.

The flowers mature sequentially from the base of the inflorescence towards the
apex. The apex is always curved over to more or less horizontal and is often covered
by the bract which subtends the entire inflorescence. The cells at the base of the
inflorescence continue to grow so that the inflorescence changes position as flowers
mature and the open flower is presented between the horizontal and around 45*^. The
number and size of the flowers in the inflorescence and the size and shape of the
bract varies between and sometimes within species but the size of the bract is a useful
character for the recognition of many species.

The pedicel is short, mostly up to only 5 mm long, and can elongate considerably
at maturity but only rarely to as long as 7.5 mm. The calyx consists of flve sepals that
are fused from the base of the calyx to half or two-thirds the length of the sepals. It
is actinomorphic, persistent and expands as it matures, providing a receptacle for the
operculate capsule that sits in the centre of the calyx. The young calyx is generally
oblong to obovate and becomes more campanulate at maturity. The lobes are linear to
triangular, often becoming broader and acuminate in maturity. The shape of the calyx
does not vary significantly between species.

The indumentum of the pedicel and calyx is similar to that on the bract and in
many species is the same on both structures. Hairs on the pedicel are to 1 mm long, to
1.5 mm long on the outside of the calyx and to 0.5 mm long on the inside of the calyx.
If present, the hairs on the inside of the calyx are usually strigose and found only on the
lobes of the calyx. The combination of type and densities of the hair on the outside of
the calyx is much more variable than on the inside of the calyx. Despite the variability
it can be a useftil character for species differentiation (e.g. E. tenerum and E. steemsii).

The corolla consists of a tube and five free corolla lobes in two lips. The tube
accounts for approximately two thirds to three quarters of the length of the corolla.
The smaller upper lip is of two lobes and the lower of three lobes. The upper lip is
sometimes much smaller than the lower. The colour of the corolla is variable. Specimen
labels indicate that the flowers may be white or pink, but usually the tube is white or
of unstated colour and the lobes are blue to violet. The flowers of many species have
irregular darker (purple) marks at the base of the lobes or in the upper throat.

In most species there are villous hairs on the inside of the eorolla tube and there
may be a minute pubescenee on the outside of the lobes. The villous hairs are often in a
dense band around the perimeter of the corolla tube and are usually placed somewhere
between half way and two-thirds of the way up the tube and below the stamens. The
tube is rarely glabrous or pubescent externally with short hairs inside. The presence
and type of hairs inside the corolla is a useful character for separating some species
of Epithema, while the hairs on the outside of the corolla lobes are not. Hairs inside
the corolla can be difficult to see in rehydrated material. The hairs on the outside of
the corolla can be difficult to observe and are not necessarily persistent. They are not
included in the species deseriptions.

The androecium consists of two fertile stamens and two staminodes. The
staminodes are a filament with an arrow- or serpent’s-head-shaped end. They are often

Revision of Epithema

163

slightly shorter than the filaments of the fertile stamens. The stamens and staminodes
are glabrous. The third, medial, staminode present in many Gesneriaceae is not seen
in Epithema. The anthers are apically coherent, the thecae are divaricate and apically
confluent. The stamens and staminodes are inserted on the top of a more or less square
flap of androecial tissue which is in turn inserted on the wall of the corolla. The flap
of tissue is up to 2.5 mm wide and high. The stamens are inserted on either end of the
flap and the staminodes to the inside of each of them. The upper edge of the flap of
tissue may be rounded leaving the staminodes at a slightly higher plane than the fertile
stamens, up to 0.5 mm higher. The head of the staminode is frequently adjacent to, but
not joined to, the back of the anther or the very top of the stamen filament. The style
grows through the space between the flap and the two coherent anthers. The androecial
tissue is normally inserted about half way to two-thirds of the way up the wall of
the corolla tube, but can be placed higher in the tube. In'espective of the position on
the corolla tube, the placement of the stamens in relation to the staminodes does not
change.

The nectary is comprised of one, two or three discrete lobes. In many species
there are two lobes, one on each side of the ovary. Sometimes the nectary lobe(s)
completely enclose the ovary, while in others the nectary is apparently absent. The
composition of the nectary is variable within and between species and is not a good
taxonomic character.

The ovary is superior, spherical to cylindrical and has one locule. The shape is
variable and is not taxonomically useful. The placentation is parietal with two ‘stalked’
placentae, one on each side of the ovary. The placentophores are fused in their lower
half to the ovaiy wall while the upper half supporting the placenta is free. This allows
the formation of a dehiscent operculum in fruit. The ovules are produced from, and
connected to, the placenta at the head of the placentophore by individual funicles.
The upper portion of the ovary is usually pubescent and the lower portion is glabrous.
Like the remainder of the plant, the indumentum consists of straight and/or hooked
hairs. The hairs are not always entirely persistent, but density, length and shape of the
hairs on the ovary are taxonomically useful. Unfortunately, the hairs are short (usually
0. 1-0.3 mm long) and not easily observed and, therefore, are of limited use for field
identification. The stigma is bilobed and the surface is covered in minute protrusions
or papillae of c. 0. 1 mm long. The style is slender and is inserted into the ovary in
the centre of the upper margin of the ovary. The style may be glabrous to densely
pubescent, the hairs being from 0.04-0.3 mm long and placed at the base of the style.
Rarely are they up to mid-way on the style.

The fi*uit of Epithema is a circumscissile or operculate capsule which is oblong,
obovate or sub-spherical in shape. The indumentum on the operculum is the same as
on the ovary although it may be sparser. After fertilisation the style withers and breaks
off leaving a small portion 0.3-2. 3 mm long. This remnant is persistent and seen on
the mature operculum. As the fruit matures, movement of the inflorescence brings the
fruit into a vertical position, the calyx lobes grow and open widely outwards, leaving
a basin-like structure in which sits the mature fruit. At maturity, the operculum is
lost, leaving the seeds open to the elements. The hair on the operculum is a useful

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Card. Bull. Singapore 67(1) 2015

taxonomic character, while fmit shape, size and length of the style remnant are not.

A large number of seeds is produced per capsule. The seeds are narrowly to
broadly ovoid, rarely sub-cylindrical to slightly sigmoid in shape, with acute and/or
constricted ends. They are not appendaged and they separate entirely from the funicle
by which they comiect to the placenta. They vary from light to dark brown and the
ends are darker than the main body of the seed. Colour can vai*y between capsules of
the same inflorescence and is certainly associated with stages of maturity, becoming
darker with age. Some variation in colour might also be an artefact of drying.

The surface of the seed coat of all species of Epithema is arranged into more or
less parallel lines that, when viewed under magnification, are darker than the remainder
of the seed coat. The lines are straight to spiralled in the direction of the long axis of
the seed and are between 0.02 and 0.05 mm apart. These longitudinal lines may fork
and cross-walls are often present between the longitudinal lines. The density of the
cross-walls and the forking does vai 7 , giving some seeds a sub-reticulate look. Older
descriptions place importance on the angle of the longitudinal lines, but angle of the
pattern varies considerably even between seeds from one capsule or different capsules
of one inflorescence. Scamiing electron microscope (SEM) photographs of the seed
of a small number of species of Epithema indicate that the surface of the seed is not
taxonomically useful (Michael Moeller and Frieda Christie, pers. comm.). Further
examination of the seed by SEM was not conducted in this revision. In only a few
species is the length of the seed useful.

Splash cup dispersal is the primary means of seed dispersal in Epithema. The
splash cups are fornied from the persistent calyx and seeds are dispersed directly from
the capsule, usually by raindrops. Splash dispersal is usually only effective over short
distances but it is also possible that seed may be transported longer distances on the fur
or the feet of animals, or in streams.

Materials and Methods

Approximately 1400 specimens from the following herbaria have been examined: A,
AAU, ABD, BK, BKF, BM, BO, C, CAL, CALI, CMU, COl, E, G, GH, K, KEP, KLU,
KYO, L, MSC, MO, NY, P, PDA, PNH, PSU, QBG, S, SAR, SING, TI, UPM, US and
WAG (Thiers, continuously updated). These specimens were collected between the
1820s and 2014 from Cameroon, Central African Republic, Democratic Republic of
Congo, Equatorial Guinea, Guinea, Ivory Coast, Liberia, Sierra Leone and Uganda, Sri
Lanka, India, Nepal, China, Taiwan, Myanmar, Thailand, Laos, Cambodia, Vietnam,
Malaysia, Singapore, Philippines, Indonesia, Timor Leste and the Solomon Islands.
All specimens cited have been seen unless othemise indicated. All measurements for
vegetative characters, peduncles, bracts, pedicels and calyces are from dried specimen
material. Measurements of other reproductive structures are taken from rehydrated
material. Observations on herbarium material have been augmented by observations
in the field (in Malaysia and Thailand) by both authors.

There are relatively few collections from areas such as eastern Thailand, Laos,

Revision of Epithema

165

Cambodia, Vietnam, Sulawesi, Maluku, The Sunda Islands, Timor and Timor Leste,
New Guinea and the many small islands of south-eastern Asia. In some of these areas
Epithema species may genuinely be absent or scarce due to the lack of ideal habitats
(e.g. absence of limestone) but collecting densities are known to be low in several of
these areas (Johns, 1995; Middleton, 2003; Newman et ah, 2007). Peninsular Malaysia
and other areas of Thailand are relatively well collected.

Provisional conservation assessments are made using the methodology proposed
by lUCN (2001).

The species are presented in alphabetical order as little data is available on
relationships between the species. The dots on the maps represent the distributions of
the specimens cited in this revision.

The flowering and fruiting times given are based on the data from herbarium
specimens only and do not preclude flowering and fruiting at other times of year.

Constraints

Flowers in an Epithema inflorescence mature sequentially within the inflorescence with
only one or few flowers fully expanded at any one time. In addition, the corolla is not
persistent. It is, therefore, common for each specimen or sheet of specimens to contain
no corolla material. This means that a thorough evaluation of infraspecific variability
in corolla, stigma, style and stamen characters was limited by the availability of
flowering material. When only immature flower buds were present they were dissected
to gather character states for corolla and ovary pubescence. It is acknowledged that
character states could change during the maturation process but we still found these
bud characters useful.

For a number of species there are very few available specimens. This limits our
understanding of the variability and distribution of the species.

Plant description, expedition date and/or geogi'aphy data was limited or absent
from many labels of the older specimens. For species with only small numbers of older
specimens, distributions below country level are unattainable and transient character
states such as corolla colour remain unknown.

Epithema Blume

Bijdr. 737 (1826); C.B.Clarke in A.DC. & C.DC., Monogr. Phan. 5(1); 176 (1883);
C.B.Clarke in Hook.f., FI. Brit Ind. 4; 369 (1884); King & Gamble, J. Asiat. Soc,
Bengal, Pt. 2, Nat Hist. 74(2): 783 (1909); Ridt, FI. Malay Penins. 2: 539 (1923);
Gamble, FI. Madras 2: 991 (1924); Pellegr., FI. Indo-Chine 4: 558 (1930); Kanjilal et
at. Ft Assam 3; 399 (1939); BameU, Ft Siam. 3(3): 205 (1962); Backer & Bakh.f ,
FI. Java 2: 527 (1965); W.L.Theob. & Grupe in Dassan. & Fosberg, Revis. Handb.
FI. Ceylon 3: 102 (1981); Pham-Hoang Ho, Cayco Vietnam ed. 3, 3(1): 24 (1993);
W.T.Wang et at, FI. China 18: 400 (1998); B.L.Burtt, Thai For. Bull. (Bot) 29: 93

166

Card. Bull. Singapore 67(1) 2015

(2001); Hilliard in Grierson & D.G.Long, FL Bhutan 2(3): 1328 (2001); A. Weber in
Kubitzki, Fam. Gen. Vase. PI. 7: 129 (2004). - TYPE: Epithema saxatile Blume.
Aikinia R.Br. in Wall., PI. Asiat. Rar. 3: 65 (1832). - TYPE: Aikinia bnmonis Wall. (=
Epithema brunonis (Wall.) Decne.).

Carpocalymna Zipp., Alg. Konst- Lett.-Bode 1: 297 (1829), nom. nud.

Herb, caulescent or acaulescent, occasionally only one leaf developing. Leaves', lowest
leaf petiolate and solitary, upper leaves petiolate or sessile, opposite or alternate; blades
thinly to thickly membranous, usually ovate to cordate, more rarely elliptic to orbicular,
if asymmetrical then the wider side up to 1 .75 times as wide as the nan'ower side, apex
rounded to acute, base mostly cordate or sub-auriculate to obtuse, inserted evenly or
unevenly on petiole, margin entire to crenate, serrate or (bi-)dentate, venation pinnate,
upper surface pale to black-green, sometimes variegated, lower surface light to mid-
or olive-green or purplish. Inflorescences l-15(-many) per plant; peduncles usually
originating from the leaf axils, occasionally from the petiole and/or the midrib of the
blade; singular bract subtending each inflorescence, cucullate and enclosing the entire
inflorescence to reduced, margin entire to dentate. Calyx cylindrical to campanulate,
consisting of a tube and 5 lobes, with an embedded gland towards the apex of each
lobe. Corolla tube usually white, lobes pale pink to blue or purple, connnonly with
darker markings on either lip; tube cylindrical to narrowly fluted, occasionally slightly
constricted at the apex; lobes with margins entire to fimbriate. Stamens and staminodes
inserted on top of flap of androecial tissue which is inserted in corolla tube; fertile
stamens 2; anthers coherent at thecae tips or along entire thecae; staminodes 2, third
staminode absent. Nectary apparently absent or one- to three-lobed, margin entire to
undulate. Ovary cylindrical to spherical, glabrous to densely pubescent, unilocular,
placentation parietal; style short; stigma bi-lobed, papillate, glabrous. Fruit (sub-)
cylindrical to (sub-)spherical; operculum circumscissile or irregularly dehiscing at
maturity, indumentum as ovary; surrounded by persistent calyx. Seed usually naiTowly
ovoid to broadly ovoid, ends acute and/or constricted, light to dark brown with darker
ends, pattern straight to spiralled, walls of pattern usually splitting and merging and
with cross- walls, occasionally walls of pattern appear thickened and rigid.

Twenty species from central tropical Africa, India, Sri Lanka, Nepal, southern China
and through Southeast Asia and Malesia to the Solomon Islands.

Key to Epithema species

la. First leaf arising more or less at ground level or from a very short stem 2

lb. First leaf arising above a definite stem 10

2a. Indumentum on ovary/operculum straight or primarily straight 3

2b. Indumentum on ovary/operculum hooked or primarily hooked, operculum
rarely glabrous 8

Revision of Epithema

167

3a. Ovary/operculum indumentum mostly villous with hairs to 1.5 mm long;
corolla > 9 mm long; Africa 20. E. tenue p.p.

3b. Ovary/operculum indumentum generally not villous, < 1 mm long (usually
much less); corolla variable; Asia 4

4a. Hairs on ovary/operculum both straight and hooked; Solomon Islands

14. E. rennellense p.p.

4b. Hairs on ovary/ operculum straight only; not Solomon Islands 5

5a. Leaf margin entire or only minutely toothed, more rarely with larger and
irregular teeth; Eastern Indonesia 1. E. longipetiolatum p.p.

5b. Leaf margin distinctly toothed; widespread in Malesia 6

6a. Ovary/operculum hairs to 0.6(-l) mm long; upper leaf surface with only
strigose to villous hairs with at least some hairs > 1 mm long; Philippines and
eastern Malesia 1. benthamii p.p.

6b. Ovary/operculum hairs to 0.3 mm long; upper leaf surface sometimes
with minute hair covering as well as longer hairs but all hairs < 1 mm long;
Philippines and Sumatra 7

7a. Leaves strongly crenate or with large, wide teeth with rounded apices; bracts >
10 mm long; Sumatra 18. E. strigosum

7b. Leaves dentate/serrate, teeth narrow with pointed apices; bracts <10 mm long;
Philippines 12. E. philippinum

8a. Hairs on ovary/operculum hooked or hooked and straight; corolla 10.4-15 mm
long; Solomon Islands 15. rennellense p.p.

8b. Hairs on ovary/operculum only hooked or operculum rarely glabrous; corolla
5-8.5 mm long; not Solomon Islands 9

9a. Inflorescences 1-5 per plant, usually large and not in fascicles; peduncles
(2-)9-27 cm long; bracts 10-25 x 6-14 mm; operculum rarely glabrous;
Sabah 4. E. dolichopodum

9b. Inflorescences 1 to many, small, often different lengths, in fascicles; peduncles
up to 10(-17) cm long; bracts 3-16(-20) x 3-10 mm; operculum with hooked

hairs; continental Asia, Taiwan, Philippines

3. E. ceylanicum (rare acaulescent forai)

10a. Ovary/operculum glabrous 11

10b. Ovary/operculum pubescent or at least with few hairs 14

lla. Plants with only 1 leaf, occasionally with 2 leaves, these never opposite;
inflorescences arising from leaf blade midrib or junction of petiole and midrib
5. E. horsfieldii p.p.

llb. Plants usually with more than 2 leaves, leaves alternate or opposite;

inflorescences in the leaf axils or opposite a solitary leaf 12

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Card. Bull. Singapore 67(1) 2015

12a. Upper leaves sessile 9. E. madulidii p.p.

12b. Upper leaves petiolate 13

13a. Leaves all or primarily alternate; 1-5 peduncles arising in sequence from
base of petiole; bracts large (10-32 x 6-12 mm) and usually enclosing
inflorescence; plants with strigose or setose hairs, inner calyx glabrous or with
sparse straight strigose hairs; Malesia 15. E. sarawakense p.p.

13b. Leaves opposite; peduncles arising from leaf axils; bracts small (4-8 x 4-9
mm) and not enclosing inflorescence, plants glabrous to sparsely pubescent,
inner calyx glabrous; southern India E. pusillum

14a. Plants with only 1 leaf, occasionally with 2 leaves, these never opposite;
inflorescences arising from leaf blade midrib or junction of petiole and midrib,
ovary hair hooked 5. E. horsfieldii p.p.

14b. Plants usually with more than 2 leaves, upper leaves usually opposite;
inflorescences Irom base of petioles or in the axils of upper petiolate or sessile
leaves, rarely at base of midrib in the rare case leaves are alternate, 1-5

peduncles arising in sequence from base of petiole 15

15a. Upper leaves sessile 16

15b. Upper leaves petiolate 19

16a. Indumentum on ovary/operculum stiuight or primarily straight; Africa

20. E. tenue p.p.

16b. Indumentum on ovary/operculum hooked or primarily hooked; not Africa

17

17a. Corolla glabrous inside; Philippines 9. E. madulidii p.p.

17b. Corolla pubescent inside 18

18a. Bract enclosing inflorescence, 13-35 x 8-12 mm 16. saxatile p.p.

18b. Bract not enclosing inflorescence, 4-16 x 4-9 mm 3. E. ceylanicum p.p.

19a. Indumentum on ovary/operculum straight or primarily straight 20

19b. Indumentum on ovary/operculum hooked or primarily hooked 24

20a. Indumentum on ovary to 1.5 mm long; Africa 2Q. E. tenue p.p.

20b. Indumentum on ovary < I mm long; Asia 21

21a. Bracts cucullate, (2-)5-23 x ( 3 -) 4 -ii mm; Philippines, eastern Malesia ... 22
21b. Bracts not cucullate, 1-13 x 2-9 mm; continental Asia and Peninsular
Malaysia 23

Revision of Epithema

169

22a. Leaves mostly not opposite, only occasionally so and then often of markedly
different sizes; leaf margin entire or only minutely toothed, more rarely with
larger and irregular teeth 1. E. longipetiolatum p.p.

22b. Upper leaves usually opposite, only rarely upper leaves solitary; leaf margin
distinctly and usually regularly toothed E. benthamii p.p.

23a. Bract small, rarely sub-cucullate or trullate, not enclosing inflorescence, 1-7
X 2-7 mm, margin often deeply lobed with 3 lobes; plants often with dense
white hairs on most plant parts; corolla 4.2-5 .4 mm long; Peninsular Malaysia
11.^. pandbracteatum p.p.

23b. Bract small or sub-cucullate, enclosing a small portion of the inflorescence,
1-13 X 3-9 mm, margin entire to dentate; plants sub-glabrous to pubescent;

corolla 5. 5-7. 5 mm long; Myanmar, China, India, Laos, Nepal, Thailand

2. E. carnosum

24a. Corolla glabrous inside 6. E. involucratum

24b. Corolla with hair (villous) inside 25

25a. Leaves all or primarily alternate; 1-5 peduncles arising in sequence from
base of petiole; bracts large (10-32 x 6-12 mm) and usually enclosing
inflorescence; Malesia 15. E. sarawakense p.p.

25b. Leaves primarily opposite; number and position of peduncles variable; bracts
small to large and enclosing inflorescence or not; Continental Asia and Malesia
26

26a. Corolla >10 mm long 8. E. longitubum

26b. Corolla <10 mm long 27

27a. Peduncles arising along petiole, often in fascicles, often many and of very
different lengths 3. E. ceylanicum p.p.

27b. Peduncles primarily arising from axils of leaves, usually singly or few and then
of not remarkably different length 28

28a. Bract only rarely sub-cucullate or trullate but not enclosing the inforescence,
1-11 X 2-12 mm 29

28b. Bract sub-cucullate to cucullate enclosing a portion or all of the inflorescence,
(3-)6-23 X 3-12 mm 31

29a. Lower to mid-style pubescent; calyx without hispid white hairs; delicate
growth forni 10. E. mej7tbranaceum

29b. Style glabrous or with only a few hairs; calyx indumentum variable but with
hispid white hairs; often with a robust growth form 30

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Card. Bull. Singapore 67(1) 2015

30a. Plants with dense white hairs on most plant parts; Peninsular Malaysia

ll. E. pai^ibracteatum p.p.

30b. Plants not densely pubescent; not in Peninsular Malaysia

3. ceylanicum (rare form with few inflorescences and smaller bracts)

31a. Bract usually enclosing only a portion of the inflorescence 32

31b. Bract usually enclosing most or all of inflorescence; corolla 6. 5-9. 5 mm long
33

32a. Plant 5-14(-20) cm; leaves often broadly lanceolate to narrowly ovate; corolla
5.2-6 mm long; Sulawesi 19. E. tenerum

32b. Plant 4-40 cm; leaves usually cordate, ovate or elliptic; corolla 5-8.5 mm
long; not in Sulawesi 3. E. ceylanicum p.p.

33a. Calyx with strigose hairs only outside, hairs 0.5-0. 8 mm long; Aceh

17. E. steenisii

33b. Calyx often glabrous, if hairs present these < 0.5 mm long; Myanmar,
Thailand, Malaysia, Sumatra, Java, Borneo, Sulawesi \6. E. saxatile p.p.

l.Epithema benthamii C.B. Clarke in A.DC. & C.DC., Monogr. Phan. 5(1): 1 80 (1 883).
- TYPE: Philippines, Albay, 1841, Cuming, H. 1265 (lectotype K [K000438696],
designated here; isolectotype BM, K [K000438697]). (Fig. 1, 2)

Epithema brunonis var. scabridum C.B.Clarke in A.DC. & C.DC., Monogr. Phan.
5(1): 178 (1883). - TYPE: Indonesia, Sulawesi, Gorontalo, Riedel s.n. (holotype K).

Epithema calcicola Ohwi, Bot. Mag., Tokyo 57: 125 (1943). - TYPE: Indonesia,
West Papua, Waren, 60 miles south of Manokwari, 26 March 1940, Kanehira, R. &
Hatusima, S. 13143 (lectotype FU, image seen, designated here; isolectotype BO).

Herb 5-30 cm high, caulescent, occasionally appearing acaulescent with leaves at
near ground level, sparsely to densely strigose and/or setose with hairs to 0.8 mm
long, the lower bract, pedicel and the outside of the calyx also minutely pubescent with
hairs to 0.2 mm long; stem 1-4 mm wide with 2-3 nodes, intemodes 0.5-6. 5 cm long.
Leaves strongly membranous, upper leaves petiolate and opposite, infrequently with a
solitary upper leaf; petiole of lowest leaf 2-1 3.5 cm long, upper petioles 0.2-6 cm long;
blade of lowest leaf 4.3-14.5 x 2.2-7. 5 cm; blades of upper leaves 2-12 x 1-6 cm,
all leaves linear, narrowly ovate, or almost oblong, occasionally ovate or rhomboid,
usually symmetrical, apex acute to rounded, base sub-cordate, rounded to truncate,
occasionally sub-auriculate, usually inserted evenly on the petiole, margin usually
in-egularly serrate or biserrate; upper surface mid-green to black-green, sparsely to
densely weakly strigose to villous, without minute hairs interspersed, straight hairs to
1 .5 mm long, hooked hairs 0.5 mm long; lower surface generally lighter in colour than

Revision of Epithema

171

Fig. 1. Epithema benthamii C.B. Clarke. A. Habit. B. Leaf. C. Infloreseenee. D. Calyx. E.
Flower opened out. F, Stigma lateral view. G. Fruit showing seeds, placenta and operculum. H.
Seeds. Scale bars: A = 6 cm, B = 3 cm, C = 5 mm, D = 3 mm, E = 4 mm, F = 1 rmu, G = 2 mm,
H = 0.5 mm. Drawn by Claire Banks from Wenzel 437 (A, C-H) and Coode 5313 (B).

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Card. Bull. Singapore 67(1) 2015

Fig. 2. Distribution of Epithema benthamii C.B. Clarke (•).

upper surface. Inflorescences l^(-6) per plant; peduncles 2.5-16 cm long, arising
from leaf axils or, rarely, from the base of the uppermost petioles; bracts sub-cucullate
or cucullate, enclosing a proportion of the inflorescence, occasionally reduced, (2-)7-
23 X (3-) 5-1 1 mm, margin usually dentate with large teeth but entire if reduced, lower
surface with long strigose hairs 0.6-1 mm long, upper surface glabrous to pubescent
at upper margin with hairs to 0.5 mm long; pedicels 1 .5^ mm long. Calyx 2.4-6 mm
long; tube 1.7-3 x 1.5-2. 5 mm; lobes 1.7-3 x 0.8-1. 5 mm; glabrous inside, rarely
with few straight hairs to 0. 1 mm long in lobes. Corolla often white, but also blue to
purple, 5-10 nmi long; tube (4-)5-6(-8) x 1.3-2. 5 mm; lobes 1-2 mm long. Stamens
1.4-1. 5 mm long; filaments 1 mm long; anthers 0.4-0.5 mm long; staminodes 0.8-0. 9
mm long. Nectary apparently absent or of two lobes, 0.8-1. 4 x 0.6-1. 1 mm, margin
entire or occasionally notched. Ovary sub-spherical to spherical, 0.9-1 x 0.9 mm,
densely pubescent to pilose, hairs straight or primarily straight with some hooked
hairs (hooked hairs are shorter and often most easily visible on the fruit operculum),
0.1-0.6(-l) mm long, on very upper portion of ovary; style 2. 9-4.2 imn long, usually
glabrous, occasionally 1 or 2 hooked hairs on base of style, 0.2 mm long; stigma 0.4
mm wide. Fruit cylindrical, ovate or obovate, 2, 1-2.3 x 1.7-1. 9 mm; operculum 0.6
mm long, pubescence as on ovary. Seed narrowly ovoid to ovoid, 0. 3-0.5 x 0. 1-0.2
mm, medium brown, pattern spiralled and reasonably even.

Distribution. Indonesia (Sulawesi, Maluku (Bum, Seram), West Papua), Philippines.

Habitat and ecology. Lithophytic on both limestone and other rocks, but also
occasionally terrestrial. Frequently found on stones and boulders in or beside streams

Revision of Epithema

173

and rivers, but also in rock crevices in humid, shady, if not wet, places. Collected from
0-1600 m altitude.

Provisional lUCN conservation assessment. Least Concern (LC). This species is
coimnon and very widespread. However, as it frequently occurs on karst limestone,
which is often commercially exploited, its status should be monitored.

Additional specimens examined: INDONESIA: Maluku: Burn, NW Bum, Bara, Waeduna
River, 30 Nov 1984, Nooteboom, H.P 5357 (L); Bum, Nov 1984, Sukendar SK740 (BO);
Maluku, Bum, Ehoe, 27 Sep 1921, Toxopeus, F.J. 358 (BO); Bum, Waeha, 12 Nov 1984, van
Balgooy, M.M.J. 4712 (E, L); Halmahera, Kampong Goal, Idjan 175 (BO); ibidem, 14 Sep
1951, Pleyte, D.R. 150 (BO, L); Seram, Tehoru, Saunulu, Murkele Ridge, 19 Feb 1985, Kato,
M. etal. C.6908 (L); Seram, Roho, 21 Nov 1917, Kornassi 482 (BO). Papua: Chaban, 28 Feb
1940, Kanehira, R. & Hatiisima, S. 11848 (A, K). Papua Barat: Raja Ampat, Misool, Near
Waima, 26 Sep 1948, Pleyte, D.R. 1053 (BO); Vogelkop Peninsula, Aitinjoe, 11 May 1958,
Versteegh, C. BW7405 (L); Biak to Parieri, 11 Sep 1966, Kostermans, A.J.G.H. & Soegeng
914 (BO, L); Biak, Kampong Saba, 31 May 1959, de Wilde, JJ.F.E. 1155 (L); Biak, 29 Aug
1915, Feuilletau de Bruyn, W.K.H. 335 (BO); Manokwari, Arfak Mts, Minjambau, 22 May
1962, Versteegh, C. BW 12694 (BO, L); Manokwari, Arfak Mts, Mupi Dessa, 16 Apr 1995,
Sands, M.J.S. 6841 (E, K); Manokwari, Arfak Plains, 25 Apr 1994, Sands, MJ.S. 6337 (E,
K); ibidem, 24 Apr 1994, Sands, M.J.S. 6318 (K); Manokwari, Kebar Valley, Andjai Airstrip,
Sobor Hills, 15 Nov 1954, van Royen, P. 5063 (A, E, L); Manokwari, Kebar Valley, Andjai
Village, 5 Nov 1 954, van Royen, P. 4946 (A, K, E); Manokwari, Mt Krabo, 3 Nov 1 960, Foster,
C. BW10773 (A, L); Manokwari, Warmare, Mokwam Trail, 10 Apr 1994, Sands, M.J.S. 6141
(K); Sorong, Warsamson Valley, 17 Aug 1961, Schram, FA.W BW 13259 (L); Wandammen
Peninsula, Wondiwoi Mountains, 28 Feb 1962, Foster, C. BW13665 (L); ibidem, 24 Feb 1962,
Schram, FA.W. BW 10631 (L). Sulawesi: s.l., 1871, RiedeH.w (P). Sulawesi Tengah: Gunung
Nokilalald, 5 May 1975, Meijer, W. 10045 (BO, L, US); Donggala, c. 57 km north of Palu on
Palu-Oti Road, 3 Jun 1979, van Balgooy, M.M.J. 3600 (L). Gorontalo: 9 Apr 2002, Mendum,
M. et al. 43 (E); Pinogu to Tulabolu, 15 Apr 2002, Mendum, M. et al. 113 (BO, E); Gunimg
Boliohutu, Mendum, M. et al. 180 (BO). Sulawesi Tenggara: Kesali-Porema, 23 Oct 1929,
Fjellbetg, G.F. 2616 (BO, S). Sulawesi Utara: G. Klabat, Pinili, 31 Oct 1973, Dransfield, J.
3869 (BO); Sangihe Island, Tamako, Kentuliang, 23 Nov 1998, Hicks, D. 151 (K).
PHILIPPINES: Agusan el Norte: Cabadbaran, Mt Urdaneta, Aug 1912, Elmer, A.D.E. 13470
(BO, E, G, GH, K, L, NY, P, US). Bohol: Aug 1923-Oct 1923, Ramos, M. BS43313 (BO, G,
K, P, US). Bukidnon: Jul 1913-Aug 1913, Escritor, L. BS21452 (US); Mt Galintan, Ramos,
M. & Edaho, G. BS48897 (NY). Cagayan: Penablanca, Mar 1929-May 1929, Ramos, M.
BS76791 (GH, SING). Davao del Sur: Sep 1911-Oct 1911, Weber, C.M. 1440 (E, NY, P).
Davao Oriental: Baguan River, 8 Mar 1949, Edam, G. 758 (A, PNH); ibidem, 8 Mar 1949,
Edaho, G. 746 (PNH); Davao, Mati, Mar 1927-Apr 1927, Ramos, M. & Edaho, G. BS49227
(NY); Davao, Mount Bilbogan, 1949, Edaho, G. 1965 (A, L, PNH). Davao del Norte: Mt
McKinley, 14 Aug \946, Edaho, G. 720 (PNH). Ifugao: Feb \9U, McGregor, R.C. BS19951
(US). Isabela: Barangay San Jose, 7 Mar 1997, Argent, G.C.G. et al. 29038 (E). Laguna:
Los Banos, Apr 1906, Elmer. A.D.E. 8318 (BO, E, G, K, NY); ibidem, 18 Dec 1903, Hallier,
H.G. 4128 (L); ibidem, 7 Mar 1906, Merrill, E.D. 5106 (BO, K, NY, US); ibidem, 14 Jan
1905, Williams, R.S. 2046 (NY); Mt Makiling, Jun 1917-Jul 1917, Elmer, A.D.E. 18283 (BO,
GH, K, US); ibidem, Jun 1917-Jul 1917, Elmer, A.D.E. 17551 (BO, GH); ibidem, Jan 1913,

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Card. Bull. Singapore 67(1) 2015

Robinson, C.B. BS17216 (US). Lanao del Sur: Lake Lanao, Cajnp Keitliley, Sep 1906-Oct
1906, Clemens, M.S. 724 (US). Leyte: Wenzel, C.A. 231 (G, US); ibidem, 28 Jun 1913, Wenzel,
C.A. 437 (GH); Mt Abucayan, Feb 1923, Edano, G. BS41748 \g, K, US). Negros Oriental:
Cuemos de Negros, Guinsuan Creek, 6 Jim 1948, Edano, G. 435a (A); ibidem, 6 Jun 1948,
Edano, G. 435 (PNH); Dumaguete, Cuernos Mts, Apr 1908, Elmer, A.D.E. 9698 (BO, E, G, L,
NY, US); Lake Balinsasayo, 1 6 Sep 1 948, Edano, G. 418 (A, PNH). Oriental Mindoro: Subaan
River, 14 Apr 1986, Coode, M.JE. 5313 (K, L). Palawan: Mt. Kabangaan, Apr 1929, Edano,
G. s.n. (GH). Quezon: Mt Cristobel, 14 Dec 1996, Reynoso & Majaducon PPI27265 (K).
Rizal: Morong, 25 Mar 1893, Loher, A. 4231 (K); Mt Angilog, Feb 1923, Lopez, G. BS42049
(BO, L, P, US); ibidem, Feb 1923, Lopez, G. BS42031 (US); Mt. Lumutan, Apr 1923, Ramos,
M. 42176 (BO). Samar: Matuguinao, Baruz, 20 Jan 1952, Gachalian, F. 147 (A, PNH); Mt
Purog, 24 Dec 1951, Edano, G. 34 78 (A); Mt Sarawag, 3 Dec 1951, Edano, G. 307 OA (PNH).
Sorsogon: Irosin, Mt Bulusan, Apr 1916, Elmer, A.D.E. 15877 (BO, C, G, GH, K, L, NY, P,
US); ibidem, 1915, Elmer, A.D.E. 15626 (BO, GH, NY). Zamboanga: Nov 1911-Dec 1911,
Merrill, E.D. 8222 (US); Sax River, 18 Feb 1905, Williams, R.S. 2401 (NY).

Notes. There are two informal forms of Epithema benthamii. The typical form is found
throughout the distribution of the species, while a finer, more thinly membranous and
more elongate fonn is found on Bum Island (Maluku) and Biak Island off Papua (e.g.
Sands 6318). These plants are similar to Epithema longipetiolatum, but still have the
typical leaf form and the characteristic long, straight or primarily straight, hairs on the
ovary and operculum of E. benthamii. The distributions of Epithema benthamii and E.
longipetiolatum overlap on Bum Island and Biak Island. See further discussion under
Epithema longipetiolatum. The indumentum of the leaf is useful for identification of
Epithema benthamii in that there are no minute hairs interspersed between the larger
hairs. The strigose hairs are often white, clearly visible and distinctive on the surface
of the dried leaves.

The specimen Kanehira and Hatusima 11848 (A) is unusual in that the peduncle
is branched and racemosely bears many complete inflorescences.

In the type specimen of Epithema brunonis var. scabrida the leaves are
somewhat dentate rather than entire as stated in the protologue, the indumentum is
of the same form as Epithema benthamii, albeit slightly denser, and the hairs on the
ovary/operculum are long and straight as in E. benthamii rather than the hooked hairs
of E. brunonis. It is quite clearly a synonym of Epithema benthamii.

The only apparent difference between Epithema calcicola and E. benthamii is
in the nan*ower somewhat reflexed bracts of E. calcicola. This species was known
only from the type collection and we conclude it is only a rather unusual specimen
of Epithema benthamii based on all other characters and that it occurs within the
distiibution of this species.

2. Epithema carnosum Benth., Scroph. Ind. 57 (1835); Barnett, FI. Siam. 3(3):
205 (1962), p.p.; Wang et ah, FI. China 18: 400 (1998); B.L.Burtt, Thai For. Bull.
(Bot.) 29: 93 (2001). - TYPE: Nepal, Nag Argute, 1832, Wallich, N. s.n. (lectotype K
[K000438688], designated here). (Fig. 3)

Revision of Epithema

175

Fig. 3. Distribution of Epithema carnosum Benth. (■), Epithema ceylanicum Gardner
(A) md Epithema piisillum (C.B. Clarke) Bransgrove (+).

Herb 6-38 cm high, caulescent; stem 1.5-5 mm wide with 2-3 nodes, intemodes
3-1 5 cm long, usually sub-glabrous to strigose or pubescent, rarely densely so. Leaves
thinly membranous to membranous, lower leaf petiolate, upper leaves petiolate or
sessile; petioles of lowest leaf 5-7.5 cm long, upper petioles 0-7 cm long; blades of
the lowest leaf 6.5-17 x 6-12.5 cm, upper leaves 2.5-12.5 x 2.3-6 cm, sub-spherical,
cordate or often ovate, symmetrical or barely asymmetrical, apex acute to broadly
acute, occasionally rounded, base sub-cordate to truncate, inserted evenly on petiole
or not, margin often sub-entire, but also serrate, dentate or crenate-sinuate (especially
upper leaves); hairs on upper surface to 1 .2 mm long; lower surface whitish-green or
paler than upper surface. Inflorescences 1-12 per plant; peduncles 0.5-7 cm long,
arising from leaf axils, from the petioles or from the leaf midribs, primarily from
the upper leaves; bracts small or sub-cucullate, enclosing a small portion of the
inflorescence, 1-13 x 3-9 mm, margin roughly entire with few dentate teeth to more
dentate; pedicels 1-5 mm long. Calyx light or whitish-green, 3. 3-5. 4 mm long; tube
2-3.3 X 1.5-2. 3 mm; lobes 1. 3-2.1 x 0.7—1 mm; indumentum outside a minute hooked
pubescence under strigose hairs 0.25-0.5 imn long. Corolla blue to dark puiple, some
with purple spots on the lower lip, 5. 5-7. 5 mm long; tube (3-)4-5.5 x (i_)i.5_2 mm
long; lobes 1-2.3 mm long; glabrous outside, villous band of hair inside. Stamen
filaments 0.6-0.8 mm long; anther dimensions unknown; staminodes 0.6-0. 8 mm
long. ISectary absent, of one lobe almost encircling ovary, or of two lobes, undulate,
each end a distinctly different height from the other, 0.6-1 x 1 .1-1.8 mm. Ovary sub-
spherical, 0.5-0.83 X 0.4-0.75 mm, pubescent or pilose, medium to high density, hairs
straight or primarily straight, hyaline, usually 0. 1-0.25 mm long, occasionally to 0.4

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Card. Bull. Singapore 67(1) 2015

mm long, on upper area of ovary; style c. 4 mm long, glabrous to pubescent, hairs
sparse to high density, to one third or half way up style, hairs hooked, hyaline, to
0.1 mm long; stigma c. 0.5 mm wide, Fmit sub-cylindrical 2. 1-2.5 x 1.7-2. 5 mm;
operculum c. 0.6 mm long, indumentum as ovary. Seed oval to fusiform, ends acute to
constricted, 0.4-0. 6 x 0. 1-0.2 mm, medium brown, pattern often straight to more or
less straight, but also to spiralled.

Distribution. India, Nepal, China, Myanmar, Thailand, Laos, Vietnam.

Habitat and ecology. Lithophytic. Found in humid, shaded places, often near water and
in rocky crevices at 0-2120 m altitude. Flowering and fruiting April, July-October.

Provisional lUCN conseryation assessment. Least Concern (LC). This species is
very widespread. However, as it frequently occurs on karst limestone, which is often
commercially exploited, its status should be monitored.

Additional specimens examined. INDIA: Andra Pradesh: Chintapalli, 31 Aug 1966, Subba
Rao, G. V. 28210 (E); Godavari, Rampa Country, Devarakonda, 6 Oct 1920, Narayanaswami,
M.A. 478 (CAL); Visakhapatnam, Araku, 24 Aug 1960, Balakrishnan, N.P. 10815 (BSI, CAL,
E). Meghalaya: Khasia, 28 Oct 1872, Clarke, C.B. 19076 (US); Khasia, Cherrapunjee, 26
Aug 1850, Hooker, J.D. & Thomson, T. 2161 (K); Khasia, Sohra, 22 Oct 1871, Clarke, C.B.
15665 (L); Mont. Khasia, Hooker, J.D. & Thomson, T. s.n. (C, CAL, E, GH, L, NY, P). Orissa:
Koraput, 8 Oct 1 950, Mooney, H.F 4045 (K); Koraput, Pottangi, 1 0 Jul 1 950, Mooney, H.F. 3853

(K) ; ibidem, 13 Jul 1950, Mooney, H.F. 3891 (K); Koraput, TuriaKonda, 10 Oct 1950, Mooney,
H.F. 4092 (K); Sikkim, 1878, King, G. s.n. (CAL); ibidem, Thomson, T. s.n. (BO, L); ibidem,
Unknown 3847 (K); Sikkim, Mongpo, 5 Oct 1884, Clarke, C.B. 36192 (G, K). Uttaranchal:
Kumaon, 25 Sep 1 884, Duthie, J.F. 2990 (K). West Bengal: Darjeeling, Unknown 12200 (BO);
Darjeeling, Mungpoo, 23 Sep 1875, Clarke, C.B. 24805 (K); Darjeeling, Rishap, 2 Aug 1870,
Clarke, C.B. 12300 (CAL, L); Darjeeling, Rishap, Rishap Ravine, 4 Sep 1869, Clarke, C.B.
8997 (L).

CHINA: Heou-hay tse, 7 Jul 1907, Esquirol, J. 711 (E). Guangxi: Tan-Ngar, 12 Jul 1928,
Clung, R.C. 6421 (A). Yunnan: Shweli-Salwin Divide, 1917-1919, Forrest, G. 15699 (E, K);
Simao, Henry, A. 13497 (K, US); ibidem, Henry, A. 12280 (K); ibidem, Hemy, .4. 12705 (K).
MYANMAR: Pegu, Scott 3 (L); Mandalay, Merktila, Taungbaw-yo, 22 Oct 1936, Smith, H.C.
16273 (K); Mandalay, Myingyan, Pope Hill, 4 Aug 1909, Lace, J.H. 4892 (E, K).
THAILAND: Chiang Mai: Chiang Dao, Doi Cliiang Dao, 22 Jul 1989, Maxwell, J.F. 89-920

(L) ; ibidem, 27 Sep 1971, Murata, G. et al. 15146 p.p. (C, K, KYO); ibidem, 27 Sep 1971,
Murata, G. et al. T.15156 (KYO); ibidem, 19 Oct 1926, Put 410 (K); ibidem, 26 Oct 1979,
Shimizu, T. et al T.20909 (KYO). Mae Hong Son: Muang, Doi Mae Sakut, 23 Sep 1995,
Nanakorn, W. et al. 4664 (E). Nan: Sapan Waterfall, 7 Sep 1995, Larsen, K. et al. 46161
(AAU). Tak: Nam Tok Phacharoen National Park, 23 Apr 2004, Poorna, R. et al. 4580 (BKF).
LAOS: Sayabourj^: Phiang, Nam Phon, 1 Sep 1999, Maxwell, J.F. 99-278 (A, L).

VIETNAM: Bac Kan: Na Ri District, Liem Thuy Municipality, Na Bo Village, 15 Jul 2004,
Atha, D. 4785 (NY); Ha Giang: Pho Bang, Pho Cao, 12 Aug 1977, Phuong, V.X. 311 (HN).

Notes. Epithema carnosum is widespread across tropical Asia with the exception of
Malesia. It is similar to some of the forms of Epithema ceylanicum and was originally

Revision of Epithema

111

separated from it (as E. dentatum) by E. carnosim having crenate-sinuate leaves vs. E.
dentatum having small teeth on the leaf margin that are sometimes subacute or doubly
seiTate. Also Epithema dentatum was said to have shorter petioles on the upper leaves.
These characters cannot consistently be used to separate Epithema carnosum from
the now more broadly defined E. ceylanicum which includes E. dentatum. It is better
separated from Epithema ceylanicum by the hairs on the ovary and operculum. They
are straight in E. carnosum and hooked or primarily so in E. ceylanicum. In nearly
all specimens from India, Nepal and Myanmar there is no variation in this character,
but on specimens from China, Thailand and Vietnam a proportion of the hairs on
the ovary/operculum of Epithema carnosum may be hooked. In all cases where the
majority of the hairs are straight, the specimens are assigned to E. carnosum. This is
also essentially tine of Epithema ceylanicum where the hairs on almost all specimens
from India and Myanmar have only hooked hairs on the ovary and operculum but
some specimens from north and north-west Thailand also have some straight hairs
on the ovai^/operculum. In all cases where the majority of the hairs are hooked, the
specimens are assigned to E. ceylanicum.

Epithema carnosum occasionally produces peduncles from the petioles of the
leaves of the tenninal leaf pair. Peduncles arising from the midrib of these leaves has
only been observed in specimens of Epithema carnosum from Khasia in Northeast
India.

3. Epithema ceylanicum Gardner, Calcutta J. Nat. Hist 6: 492 (1846). - Epithema
carnosum var. ceylanicum (Gardner) C.B. Clarke in A.DC. & C.DC. Monogr. Phan.
5(1): 178 (1883). - TYPE: Sri Lanka, Gardner, G. 606 (lectotype K, designated here).
(Fig. 3-5)

Epithema carnosum var. hispidum C.B. Clarke in A.DC. & C.DC., Monogr. Phan.
5(1): 178 (1883), - Epithema dentatum subsp. hispidum (C.B. Clarke) Hilliard &

B. L.Burtt, Edinburgh J. Bot. 54: 112 (1997). - TYPE: India, Tamil Nadu, Western
Ghats, Courtallum, August 1 835, Wight, R. 2350 (lectotype K, designated here)

Epithema carnosum var. dentatum C.B.Clarke in A.DC. & C.DC., Monogr. Phan. 5(1):
111 (1883), - Epithema dentatum (C.B.Clarke) Hilliard & B.L.Burtt, Edinburgh J.
Bot. 54: 111 (1997). - TYPE: Bunua, Mon, Moulmein, Farm Cave Rocks, Parish,

C. S.P 63 (lectotype K, designated by Hilliard & Burtt (1997)).

Epithema brunonis veiv. fasciculatum C.B.Clarke in A.DC. & C.DC., Monogr. Phan.
5(1): 180 (1883). - Epithema taiwanense vm. fasciculatum (C.B.Clarke) Z.Yu Li
& M.T.Kao, FI. Taiwan ed. 2, 4: 697 (1998). - TYPE: Philippines, Tayabas, 1841,
Cuming, H. 823 (lectotype K [K000438698], designated here; isolectotype BM,
G-DC, K, L, P).

178

Card. Bull Singapore 67(1) 2015

Fig. 4. Epithema ceylanicum Gardner. Leaves and inflorescenees. (Photo: Preeeha Karaket)

Fig. 5. Epithema ceylanicum Gardner. Flowers. (Photo: Preeeha Karaket)

Revision of Epithema

179

Epithema taiwanense S.S.Ying, Coloured 111. FI. Taiwan 4: 795 (1992); Li & Kao, FI.
Taiwan ed. 2, 4: 697 (1998); Wang et al., FI. China 1 8: 400 (1998). - TYPE: Taiwan,
Chia-yi, Kuanyin waterfalls, 200-300 m, 25 June 1 992, Yirjg, S.S. s.n. (holotype NTUF
n.v.).

Herb 4^0 cm high, caulescent or with very short primary intemode and appearing
acaulescent; stem 1-10 mm wide, intemodes 0.5-15 cm long, sub-glabrous, sparsely
strigose or sparsely pubescent, hairs 0.5-1 imn long. Leaves thinly to strongly
membranous, lowest leaf always petiolate, upper leaves sessile to petiolate, sometimes
with a solitary upper, sessile leaf per plant; petiole of lowest leaf (0.2-)l-12 cm long,
sub-glabrous to densely pubescent, hairs to 0.7 mm long, petioles of upper leaves 0-2
cm long, sub-glabrous, minutely pubescent or with longer hairs of medium to high
density, to 1 mm long; blades of lowest leaves 3-20 x 1.8-20 cm, upper or solitary
leaves ( 1 -) 1 .5-1 1 x 0.8-7 cm, lower leaves cordate to ovate, upper leaves cordate, ovate
or elliptic, occasionally almost oblong or sub-spherical, symmetrical or asymmetrical,
apex acute to rounded, base of lower leaf cordate to truncate, base of upper leaves
obtuse, rounded or truncate, occasionally sub-cordate, inserted evenly on petiole or
not, where inserted unevenly one side is often cordate, the other truncate, margin sub-
entire to dentate, bidentate or seirate; upper surface pale- to mid-green, with sparse
straight and/or hooked hairs to 0.9 mm long and sparsely to densely strigose, hairs
0.25-1 .5 mm long; lower surface paler green or purplish in colour, sub-glabrous to
strigose, hairs sparse to medium density, sometimes with hooked and straight hairs on
veins to 0.3 mm long. Inflorescences 1-6 (to many) per plant; peduncles arising from
the leaf axil, the petiole or the base of the midrib, usually from the upper leaves, often
rather fasciculate, (0rt-)l-10(-17) cm long, often of very variable lengths in a single
cluster, sub-glabrous to densely setose, hairs to 1.1 mm long; bracts (sub-)cucullate
but enclosing a small proportion of the inflorescence if at all, 3-16(-20) x 3_9(_i0)
mm, 0.6-2(-3.4) times as long as wide, margin entire to partially or entirely dentate;
lower surface glabrous to strigose, hairs sparse to medium density, hairs 0.25-0.8 mm
long, with or without shorter layer of hair 0. 1-0.4 mm long; upper surface glabrous
to sparsely strigose, hairs to 0.25 nun long, hairs on upper half of bract or towards
margins; pedicels 0.5-6 mm long, densely pubescent with fine minute hairs and more
sparsely with larger strigose hairs to 1 mm long. Calyx 2.1-8 mm long; tube 0.8-6 x
1-2.9 mm; lobes 1-2.5 x 0.7-2. 1 mm; sub-glabrous or sparsely to densely pubescent
outside with fine, minute straight and hooked hairs to 0.25 mm long, with or without
sparse, strigose hairs, to 0.4 mm long; glabrous or subglabrous inside, sometimes with
strigose hafts on inside of lobes. Corolla whitish or blue, blue-violet or pink, tube
white, 5-8.5 mm long; tube 3.2-6.3 x 1-3 mm; lobes 1-3 x 1-1,3 mm long, margins
often slightly fimbriate; glabrous to sparsely pubescent outside on lobes, hairs straight;
dense band of villous hair inside, one-third to three-quarters way up the tube from the
base. Stamens 1. 4-2.1 nun long; filaments 1-1.7 mm long; anthers 0.4-0. 5 mm long;
staminodes 1-1.7 mm long. Nectary apparently absent or one or two discrete lobes,
may almost encircle ovary or not, margin entire or sub-entire, undulate or notched,
0.8-1. 8 X 1-1.8 mm. Ovary sub-spherical to cylindrical, 0.7-1. 5 x 0.9-1. 5 mm,

180

Card. Bull. Singapore 67(1) 2015

sparsely to densely pubescent on the upper half or uppermost part of the ovary, hairs
hooked or primarily hooked, hyaline, to 0.15 mm long; style 2.5-5 mm long, glabrous
or sparsely pubescent in lower half, hairs hooked; stigma to 0.5-0. 8 mm wide. Fruit
spherical to cylindrical, 1.5-5 x 1-3 mm; operculum 0.4-0. 8 mm long, sparsely to
densely pubescent as on ovary. Seed sub-cylindrical to ovoid, 0.4-0. 7 x 0. 1-0.2 mm,
pattem often straight or almost straight, but also to spiralled, regular, walls clearly
defined, often appearing thickened and rigid.

Distribution. India (including Andaman Islands), Sri Lanka, Taiwan, Myanmar,
Thailand, Cambodia, Vietnam, Philippines.

Habitat and ecology. Lithophytic. Found on rocks and rock faces in wet, humid, shady
places, occasionally on wet tree trunks at 0-2000 m altitude.

Provisional lUCN conset^ation assessment. Least Concern (LC). This species is
very widespread. However, as it frequently occurs on karst limestone, which is often
commercially exploited, its status should be monitored.

Additional specimens examined. INDIA: Andaman and Nicobar Islands: North Andaman,
Saddle Peak National Park, 10 Feb 2000, Sumathi, R. 17895 (K). Karnataka: Chorla
Ghats, Chorla, Stocks s.n. (K); Coorg, Mercara, Jersey Falls, 12 Sep 1934, Barnes, E. 928
(K); Mysore, Hassan, Bisle ghat, 14 Aug 1971, Ramamoorthy, T.P. HFP2007 (US); Mysore,
Hassan, Shiradi ghat, 7 Aug 1969, Saldanha, C.J. 14426 (US); Mysore, Kenchankumri State
Forest, 15 Aug 1971, Ramamoorthy, T.P. HFP2035 (K, US); Mysore, Shimoga, Hulical ghat,
24 Aug 1963, Raghavan, R.S. 90185 (E). Kerala: Idikki District, Sabarimalai slopes, 26 Sep
1972, Sharma, B.D. 42032 (E); Kochi (Cochin), Parambikulam, 25 Sep 1935, Barnes, E. 1300
(K); Kochi (Cochin), Parambikulam Hills, Dec 1934, Barnes, E. 1056 (K); Palghat, Silent
Valley, 21 Aug 1966, Vajvavelu, E. 27558 (L). Madhya Pradesh: Bastar, Upper Kanger Valley,
26 Aug 1959, Subramanyam, K. 8644 (E). Maharastra: Concan, Hooker, J.D. & Thomson,
T. s.n. (P); ibidem. Stocks, Law & co. s.n. (C, L). Orissa: Dhenkanal State, Goyalpathar ghati
valley, 16 Sep 1942, Mooney, H.F 2067 (K); Dhenkanal State, Kapilas Hill, Mooney, HE 2754

(K) ; Dhenkanal State, Saptasajya, 20 Sep 1993, Dhal, N.K. & Rout, N.C, 8122 (E); Ganjam,
Budhakliol Pahad, 14 Aug 1971, Brahmam, M, 2776 (E). Tamil Nadu: Banliyar, ?Wight s.n.

(L) ; ibidem, ? Wight s.n. (L); Madurai, Way, 14 Oct 1959, Subramanyam, K. 8975 (E); Mont.
Nilghiri & Kurg, Thomson, G. 121 (K); ibidem, Thomson, G. s.n. (E, G); Vilangadu Forest, 27
Aug 1985, Pradeep, A.K. 5850 (CAL); Western Ghats, Courtallum, W. C. 2680 (E); ibidem,
Wight, R. 556 (K); Western Ghats, Courtallum, Swamiar Falls, 2 Oct 1975, Nan; K.K.N. 1354
(CALI). West Bengal: Kolcata, Kabini tributory falls, 17 Aug 1964, Ellis, J.L. 20452 (E).

SRI LANKA: s.l., 1891, Deschamps s.n. (G); Mrs. General Walker 140 (K); Mrs. General
Walker s.n. (K); Thwaites, GH.K. 2844 (BO, G, K, P); 1 836, Wight, R. 1/162 (K). Uva: Badulla
District, Between Mesitale and Dunhinda Falls, 26 Jan 1982, Namba, T & Mikage, M. 82-
413 (KYO); Badulla District, Dunhinda Falls, 8 Jan 1983, Lucas 1025 (US). Central: Matale
District, Wiltshire Forest, 2 Jul 1974, Sumithraarachchi, D.B. DBS388 (K); Deltotte, Nov
1888, Unknown s.n. (SING); Kandy, Laksapana, 19 Sep \912, Jayasuriya, A.H.M. 843 (US);
Peradeniya, 10 Sep 1934, Unknown 753 (L). Sabaragamuwa: Kegalle District, Dolosbage,
Windsor Forest, 17 Nov 1978, Grey-Wilson, C. & Grey-Wilson, C.M. 3016 {K. US); Kegalle
District, Kadaganawa, 1 Sep 1969, Grupe, D.A. 200 (US); ibidem, 22 Aug 1968, Theobald,

Revision of Epithema

181

W.L. & Gmpe, D.A. 2404 (US); ibidem, 30 Jul 1968, Theobald, W.L. & Grupe, D.A. 2368 (US);
Ratnapura, Katussagala Hill, 5 Dec 1976, Faden, R.B. & Faden, A.J. 76/494 (US); Ratnapura,
Pannil Kanda, 5 Jul 1975, Waas, S. 1407 (K); Ratnapura, Rakwana, 7 Dec 1977, Fosberg, FR.
57278 (K, US).

TAIWAN: Takow, Ape’s Hill, Hemy, A. 1916 (E, K, NY, US); Kaohsiung Hsien, Shuangchi,
Meilong, 31 Oct 1988, Hsieh, C.F et al. s.n. (E); Kaohsiung Hsien, Takangshan, 14 Nov 1988,
Yang, K.C. 3465 (E).

MYANMAR; Mon: Moulmein, Jul 1880, Brandis, D. 676 (K); ibidem, Lobb, T 387 (K);
ibidem, McKee, H.S. 1958 (P); ibidem, Parish, C.S.P. 133 (K); Moulmein, Kyauk Ta Lon, Jul
1958, McKee, H.S. 6317 {K, P).

THAILAND: Bueng Kan: Mueang, Chaiyaphon, Singhanat Ban Phot Temple, 26 Aug 2001,
Pooma, R. et al 2697 (BKF). Chiang Mai: Chiang Dao, Pooma, R. 1059 (BKF); ibidem, 16
Jul 1958, Smitinand, T. 47/P (BKF); ibidem, 15 Aug 1963, Smitinand, T. & Sleumer, H.O. 1006
(BKF); Chiang Dao, Doi Chiang Dao, 27 Sep 1971, Murata, G. et cd. 15146 p.p. (L); ibidem,
17 Jul 1958, Sorensen, T et al. 4235 (A, BKF, C, E); Chiang Dao, Doi Chiang Dao, Between
Pong Pho and Khun Klong, 3 1 Jul 1 968, Larsen, K. et al. 2923 (AAU); Doi Sutep, 1 0 Oct 87,
Maxwell, J.F. 87-1154 (BKF, L); ibidem, 12 Sep 1958, Sorensen, T. et al. 4936 (ABD, C); Mae
Ram Subdistrict, Doi Sutep-Pui National Park, Mae Ram Stream, 10 Sep 1992, Palee, P. 73
(A, CMU, GH, L); Fang, Doi Ang Khang, 21 Sep 2008, Middleton, D.J. et al. 4544 (E); Mae
Dang, Mawn Ngaw, Doi Mawn Ngaw, 12 Aug 2002, Palee, P. 544 (L). Chumphon: Sapli, 8
Sep 1927, Put 1015 (ABD, K). Kanchanaburi: 15 Aug 1968, Nimanong, B. & Phusomsaeng,
S. 288 (BKF); ibidem, Suvanakoses, P. 2060 (BKF); Kanburi, Baw Re, 21 Jul 1926, Put 219
(K); Sai Yok, Sai Yok National Park, Lam Tam Keio, 10 Aug 1982, Shimizu, T. et al. T.28519
(BKF, SING); Sangklilaburi, 28 Jul 1968, Prayad 1422 (BKF); Thong Pha Phum, 4 Jul 1973,
Maxwell J.F. 73-107 (AAU, BK); Wang Pho, 26 Oct 1969, Kasem 641 (BKF); Khon Kaen,
Chum Phae, Phu Pha Man national park, Tham Pha Puang, 1 Aug 2011, Nonsaengsri, M. et
al. 7965 (QBG); Khon Kaen, Nanong Turn, Ban Na Chan, 23 Aug 2011, Norsaengsri, M. et
al. 8048 (QBG); Khon Kaen, PhaNok Khao, 9 Sep 1963, Smitinand, T & Sleumer, H.O. 1109
(K, L, SING). Lampang: Doi Pang La, Huay Tak, 25 Sep 1967, Shimizu, T. et al. T. 10789
(BKF); Muang Bahn, Chae Son, Jae Sawn Stream, 22 Aug 1995, Maxwell, J.F. 95-536 (BKT,
CMU, L); Ngao, Near Tham Pha Thai, 24 Sep 2008, Middleton, D.J. et al. 4579 (E); Loei,
Phu Kradung, 15 Oct 1967, Prayad 1034 (BKF). Loei: Phu Ruea District, Phu Luang, 16
Sep 1966, Phusomsaeng, S. & Bunchuai, K. 42 (BKF, L). Mae Hong Son: Muang, Doi Pui,
23 Sep 1995, Larsen, K. et al. 46861 (AAU); Mueang, Tliam Pla-Namtok Pha Suea National
Park, Pha Daeng Cave, 22 Aug 2013, Norsaengsn, M. 10568 (QBG). Nakhon Nayok: Nang
Rong Falls, 4 Aug 1992, Larsen, K. et al. 43771 (P). Nan: Tham Pha Toop Forest Park, 2 Sep
1999, Middleton, D.J. 149 (A, BKF, E, K); Muang, Tham Phatup Forest Park, 16 Aug 2012,
Middleton, D.J. et al. 5613 (E, SING); Muang Tam PaaToop, 13 Sep 1995, Nanakorn, W. 4240
(QBG); Tham Pha Tok, 25 Jul 1992, Larsen, K. et al 43589 (AAU); ibidem, 13 Sep 1995,
Larsen, K. et al 46426 (AAU). Phayao: Doi Luang National Park, Champa Thong Waterfall,
9 Aug 1997, Petrmitr, O. 70 (L). Prachuap Khiri Khan: Bang Saphan, Khao Maa Rong, 5 Sep
2008, Middleton, D.J. et al. 4284 (E), Phrae: Nam Pu, 23 Sep 1 9 1 2, Van Pruk, L. 350 (BK, K).
SaKaeow: KhaoChakan, 17 Oct Staples, G.W. etal. 1409 {E, SING). Saraburi: Kaeng
Khoi, Amata-khuha Cave, 20 Aug 2001 , Pooma, R. et al 2100 (BKF); Kaeng Khoi, Tham Pra
Photisat, 7 Oct 1979, Shimizu, T. et al. T. 194 13 (KYO, L); Muak Lek, 10 Nov 1924, Marcan,
A. 1869 (ABD, K); ibidem, 3 Sep 1928, Put 1853 (K); Phraphutthabat, Wat Khaowong, 18 Sep
2004, Pooma, R. et al. 4824 (E), Surat Thani: Kanchanadit, 31 Jul 1927, Kerr, A.F.G. 13053
(K); Koh Samui, Kao Noi, 28 Jun 1966, Sakol 1150 (BKF). Tak: Mae Sot, Doi Muser, 20 Aug

182

Card. Bull. Singapore 67(1) 2015

1961, Chermsirivathana, C. 1 (BKF); Mae Sot, Khao Pha Wo, 23 Jul 1973, Murata, G. et al.
T.16922 (AAU, BKF, KYO, L); ibidem, 12 Oct 1979, Shimizu, T. et al. T.18519 (KYO); Mae
Sot, Wat Tham Inthanin, 11 Sep 2009, Middleton, D.J. & Triboun, P 4846 (E). Tak: Umphang,
Takhobi Cave, 18 Oct 2014, Middleton, D.J. et al. 5768 (BKF, E, SING). Uthai Thani: Lan
Sale, Huppatat Non Flunting Area, 14 Oct 2014, Middleton, D.J. et al. 5687 (BKF, E, SING).
CAMBODIA: Kampot: 20 Sep 1903, Geoffray, M. 95 (P).

VIETNAM: Lang Son: hmg\mc,Eberhardt 3363 (P).

PHILIPPINES: Apayao: May 1917, Fenix, E. BS28106 (US). Benguet: Mt Trail, Oct 1929,
Quisumbing, E. BS78105 (NY); Twin Peaks, 8 Sep 1904, Williams, R.S. 937 (NY). Bukidnon:
Tanculan, Jul 1916, Fenix, E. BS26049 (K, NY, US). Bulacan: 8 Sep 1935, Bartlett, H.H.
14744 (GH), Ilocos Norte: Bangui, Nov 1923, McGregor, R.C. BS43566 (K); Burgos, Jul
1918, Ramos, M. BS32892 (GH, K, US). Laguna: Los Banos, Loher, A. 1555 (K). Oriental
Mindoro: Mansalay, Mt Yagaw, Nov 1952-Dec 1952, Sulit, M.D. & Conklin, H.C. 16913 (L,
PNH). Pampanga: Mt Arayat, Sep 1905, Merrill, E.D. 4216 (K). Rizal: Aug 1911, Ramos, M.
BS13624 (BO, G, K, L, P, US). Rizal: Antipolo, Jul 1917, Ramos. M. & Edaho, G. BS29466
(BO, P, US); Kay Ungulan, 18 Aug 1935, Bartlett, H.H. 15380 (GH); Montalban, 11 Aug 1935,
Bartlett, H.H. 14487 (GH, PNH); ibidem, 3 Aug 1990, Loher, A. 1556 (K); ibidem, Dec 1904,
Loher, A. 6669 (K); ibidem, Dec 1904, Loher, A. 6668 (K, US); ibidem, 26 Sep 1890, Loher, A.
1554 (K, P, US); ibidem, Nov 1909, Merrill, E.D. 52 (G, US); ibidem, 2 Nov 1909, Robinson,
C.B. BS9534 (NY); ibidem, 2 Nov 1909, Robinson, C.B. BS9525 (L, US); Morong, Aug 1906,
Ramos, M. BS1435 (NY, US).

Notes. Epithema ceylanicum is a widespread and variable species. The protologue of
Epithema ceylanicum states the plants are hispidly pilose, have a persistent style and
have five distinct nectaries. Only a proportion of the specimens from Sri Lanka are
densely pubescent, only the basal section of the style is persistent and we have seen no
more than two nectaries. As a variety Clarke (1883) separated Epithema carnosum var.
ceylanicum (as ""zeylanica'") from E. carnosum var. carnosum on its larger calyx and
seeds that are twice as large. However, neither of these characters holds up to closer
scrutiny. Instead, it is easily distinguished from Epithema carnosum by the hooked
rather than straight hairs on the operculum.

Rarely, it can be difficult to differentiate Epithema ceylanicum fromE’. saxatile,
particularly as they both have hooked hairs on the ovary/operculum. They are most
easily distinguished in the size of the bract which in Epithema saxatile usually encloses
the inflorescence.

Epithema taiwanense was previously distinguished by the presence of many
short, often fasciculate, peduncles with very small inflorescences and quite consistently
rounded or rounded-acute leaf apices. However, these characters are within the overall
range of variation of the very variable Epithema ceylanicum and we have, therefore,
decided to synonymise the names and recognise a single species.

Some specimens bear a resemblance to Epithema involucratum but they can be
distinguished by the presence of hair in the corolla and the usually numerous, small
inflorescences among other characters. One of the specimens from the Philippines,
Fenix BS28106 (US), somewhat resembles Epithema benthamii in leaf shape but is,
again, easily separated by having hooked hairs on the ovary/operculum rather than the
straight hairs of E. benthamii.

Revision of Epithema

183

One collection from the Philippines, R.S. Williams 937 (NY(x2)), is rather
distinct from most of the material of this species in the Philippines in its acaulescent
habit and sessile upper leaves. It also has large bracts. None of these characters is
unique in the species but the combmation is unusual. Further collections are necessary
to see whether this is distinct from Epithema ceylanicum.

Some collections of this species have strongly variegated leaves but this
character does not appear to have any taxonomic significance.

4. Epithema dolichopodum Hilliard & B.L.Burtt, Edinburgh J. Bot. 54: 112 (1997).
- TYPE: Malaysia, Sabah, Lahad Datu, Bukit Tempadong, 100-150 m, 11 June 1984,
Beaman, J.H. 10068 (holotype E; isotypes GH, K, L, MSC, NY, US). (Fig. 6)

Herb 7-28 cm tall, acaulescent with first leaves arising at ground level, sometimes
with 1-2 additional elongated intemodes, indumentum of vegetative organs sparsely
to densely strigose and/or setose, hairs to 0.6 mm long; stem 2-6 mm wide with
0-2 nodes, intemodes to 8 cm long. Leaves membranous to sh'ongly membranous,
petiolate, solitary, opposite one or more peduncles; petioles (1.5-)5-37 cm; blades

5- 24 X 4-15.5 cm, cordate, broadly ovate or occasionally sub-spherical, symmetrical
to barely asymmetrical, apex usually rounded, occasionally acute, base cordate or
sub-truncate, occasionally sub-auriculate, inserted evenly on petiole or not, margin
sub-entire to dentate and bidentate. Inflorescences 1-5 per plant; peduncles (2-)9-27
cm long, originating from the base of the plant or opposite solitary leaves; bracts
pale green or green, cucullate but not always enclosing entire inflorescence, 10-25 x

6- 14 mm, 1.2-2. 5 times as long as wide, margin dentate; pedicels 2. 1^.6 mm long,
sub-glabrous to densely pubescent, hairs minute and hooked. Calyx lobes purple or
purple only at tips, 3. 8-5. 6 mm long; tube 2-4 x l .5-3 mm, lobes triangular, 1 .3-2.7 x
0.8-2 mm; minutely densely pubescent outside, hairs white; glabrous to very sparsely
pubescent inside, hairs minute, in tips of lobes. Corolla tube white, lobes blue, with or
without piiiple markings on lobes, 5-8.4 mm long; tube cylindrical, 3. 3-6. 3 x 1.3-3
mm, lobes 1 ,7-2. 1 mm long, margin slightly fimbriate to fimbriate; band of villous hair
inside. Stamen filaments 1 mm long; anther characters unknown; staminodes 1 mm
long. Nectary of two discrete lobes, 0.4-1. 5 x 0.8-1 .7 mm, margin undulate. Ovary
sub-cylindrical to sub-spherical, 0.8-1. 3 x 0.6-1 .3 mm, minutely pubescent on upper
portion of ovary, medium to high density, hairs hooked, hyaline or white, 0.04-0.1 mm
long; style 3.7-4. 1 mm long, glabrous or rarely with minute, hooked hairs; stigma c.
0.5 mm wide. Fruit cylindrical to obovate, 2.3-3 x 2. 1-2.5 mm; operculum 0.6-0. 8
mm long, glabrous or as on ovary. Seed narrowly ovoid to ovoid, 0.3-0. 6 x 0. 1-0.2,
dark brown, pattern partially spiralled or spiralled, regular.

Distribution. Malaysia (Sabah), Philippines (Palawan).

Habitat and ecology. Lithophytic on limestone. Found on boulders, outcrops, cave
entrance walls in humid and shaded places at 90-150 m altitude. Flowering and
fruiting in June, September-November.

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Card. Bull. Singapore 67(1) 2015

Fig. 6. Distribution of Epithema dolichopodum Hilliard & B.L.Burtt (A), Epithema horsfieldii
(R.Br.) DC. (•), Epithema madulidii Hilliard & B.L.Burtt (★), Epithema philippinum (Hilliard
& B.L.Burtt) Bransgrove (+) and Epithema tenerum (C.B. Clarke) Hilliard & B.L.Burtt (■).

Provisional lUCN conservation assessment. Least Concern (LC). Although this species
is not especially widespread, its distribution is not restricted enough to fall into the
Vulnerable category under the B criteria if there were accompanying threats. However,
as it fi'equently occurs on karst limestone, which is often commercially exploited, its
status should be monitored.

Additional specimens examined. MALAYSIA: Borneo: Sabah: Hutan Simpan Madai,
15 Sep 1976, Tamura, M. & Hotta, M. 690 (KYO); Kudat, Pulau Balembangan, Oct 1994,
Anthonysamy, S. etal. SB7 (E); Kudat, Pulau Balembangan, Kampung Sina, 9 Apr 1977, Stone,
B.C. & Anderson, E.F SAN86719 (KLU); Kunak, Madai Hill, 9 Jun 1996, Lim, S.P et al.
LSP681 (SING); ibidem, 9 Jun 1996, Lim, S.P. et al. LSP675 (SING); Lahad Datu, Segama
River, Batu Sarang, 25 Aug 1999, Kiew, R. RK4759 (SING); Madai Baturong Forest Reserve,
Madai Caves, 14 Nov 1968, Kokawa, S. & Hotta, M. 1125 (KYO, L); ibidem, 14 Nov 1968,
Kokawa, S. & Hotta, M. 1073 (KYO); Segarong, Kiew, R. et al. RK4321 (K, SENG); ibidem,
Symington, C.F. & Agama, J. 9361 (K, SING); Sukau, Bod Tai, 14 Sep 1996, Kiew, R. & Lim,
S.P. RK4138 (K, SING); Tawao, Oct 1922-Mar 1923, Elmer, A.D.E. 20569 (G, GH, K, L,
SING); Tawau District, Mount Wullersdorf Forest Reserve, Teck Guan Quarry, Nadiah, I. et
al. SAN149136 (SING).

PHILIPPINES: Palawan: Mt Kabangaan, Apr 1929, Edano, G. BS77711 (GH); Mt
Mantalingajan, Apr 1929, Edano, G. BS77570 (GH, SING).

Notes. This species is most easily confused with other acaulescent species such as
Epithema strigosum, E. longipetio latum and^”. rennellense. One of the differentiating
features of Epithema dolichopodum are the very long peduncles which are usually

Revision of Epithema

185

(l-)2(-3) times the height of the rest of the plant. In Epithema rennellense, E.
longipetiolatum and E. strigosum the peduncles are shorter to barely longer than the
height of the plant. Other differences are discussed in the notes section of the other
species.

The hairs on the ovary and operculum are hooked and minute, the only variation
being the density of the hairs. While it is infrequent, the lower style may be pubescent
with hooked hairs. Fruit with glabrous and pubescent styles were observed from the
same plant.

While the upper surface of the leaves can be densely strigose, the surface of the
leaves and the stem may appear glabrous due to the small size of the hairs.

The collections available for examination are from relatively few sites, four of
which were in close proximity to each other.

To-date, the only other species of Epithema found in Sabah are E. saxatile and
E. sarawakense.

S. Epithema horsfieldii (R.Br.) A.DC,, Prod. 9: 279 (1845). -Aikinia horsfieldii R.Br.,
PL Asiat. Rar. 3: 66 (1 932). - TYPE: Indonesia, Java, 1 802-1818, Horsfield, T 85, also
labelled Cyrtandn 3 (lectotype BM [BMOO 1125214], designated here; isolectotype K
(x2)). (Fig. 6)

Epithema difformis Span., Linnaea 15: 331 (1841). - Carpocalymna monophylla
Zipp. ex Span., Linnaea 15: 331 (1841), nom. inval. - TYPE: Indonesia, Timor, 1841,
Spanoghe, J.B. 57 (lectotype P, designated here; isolectotype L).

Epithema horsfieldii var. epiphyllum Hilliard & B.L.Burtt, Edinburgh J. Bot. 54: 112
(1997). - TYPE: Indonesia, Western Sumbawa, Batudulang and Sampar Olat Ranges,
Mt Batulanteh, April 1961, Kostermans, A.J.G.H. 18839 (holotype L; isotypes BO
(image seen), K).

Herb 3.2-22 cm high, caulescent (but can appear as just one large leaf with the stem
mistaken for a petiole), unifoliate or rarely with two leaves; stem 0.5-3. 8 mm wide.
Leaves sessile (but see above), membranous to strongly membranous; blade 3.2-17.5
X 2.4-13 cm, usually cordate or sub-cordate but also to oblong, symmetrical or not,
apex broadly acute to rounded, base cordate to truncate, inserted evenly on petiole,
margin entire to dentate, bidentate, sen'ate or crenate, arrangement of veins variable
but the lower secondary veins often appearing to arise from around the base of the
peduncles, often widely spaced, occasionally running down sides of midrib; upper
surface pale green to green; lower surface light green or purplish. Inflorescences 1-1 5
per plant; peduncles 0.3-13.3 cm long, originating from the midrib at the base of the
blade, or from the midrib up to 2.3 cm from the base; bracts cucullate, sub-cucullate
or small and only enclosing a small proportion of the inflorescence, 2-22 x 3-12 mm,
margin variable, irregularly dentate to dentate, occasionally lobed; pedicels 0.7-5 mm
long. Calyx 2. 5-5. 7 mm long; tube 1-4.4 x 0.9-2. 7 mm; lobes 0.6-5 x 0.4-2. 3 mm;

186

Card. Bull. Singapore 67(1) 2015

glabrous inside or rarely sub-glabrous with hairs to 0.1 mm long. Corolla blue, lilac,
lilac-blue, violet-purple and white in proximal half of the two upper lobes or violet
only at tips, 6.5-12 mm long; tube cylindrical 4-9 x 0.9-2 mm; lobes 1.5^.2 x ]
mm, margin undulate to slightly fimbriate; band of villous hair inside at one third to
half way up the tube, occasionally the band is incomplete or the hair sparse. Stamens
0.7-1. 9 mm long; filaments 0,6-1. 7 mm long; anthers 0.3-0. 6 mm long; staminodes
0.4-1 mm long. Nectary apparently absent or one or two lobes, sometimes encircling
the ovary, margin entire or undulate and of variable height, 0.2-1. 3 x 0. 1-1.5 mm.
Ovary sub-cylindrical to sub-spherical, 0.7-1. 6 x 0.5-1. 7 mm, sparsely to densely
pubescent, hairs hooked to 0.1 mm long, on upper half of ovary or the uppermost
portion of the ovary below the base of the style; style 0.5-5. 8 mm long, glabrous to
pubescent at the base, hairs hooked, 0.04-0.2 mm long; stigma 0.3-0.6 mm wide.
Fruit obovate-cylindrical, shortly cylindrical or sub-spherical, 1.3-3 x 1-3 mm long;
operculum 0.5-1 mm long, glabrous or pubescent in upper parts, hairs medium to
high density, hooked, white, 0.04-0.13 mm long. Seed narrowly ovoid to ovoid,
infrequently slightly sigmoid, ends acute or constricted, 0.3-0. 6 x 0. 1-0.2 mm, light
to dark brown, pattern straight to spiralled and more or less regular.

Distribution. Indonesia (Java, Lesser Sunda Islands, Sulawesi), Timor Leste.

Habitat and ecology^. Primarily lithophytic on limestone hills, rocks and cliffs, usually
in shaded to heavily shaded areas. Also found in roadside cuttings. Collected at 40-
900 m altitude. Flowering and fruiting January-May.

Provisional lUCN consei^ation assessment. Data Deficient (DD). Although this
species is widespread and would normally be considered to be Least Concern there
are surprisingly few recent collections from much of its range, particularly from Java.
It is unclear whether this is due to a lack of collecting activity or whether it really has
declined over much of its range. This will need to be investigated before a realistic
conservation assessment can be made.

Additional specimens examined. INDONESIA: Bali: Singaraja, Gitgit Waterfall, 5 Apr 1936,
van Steenis, C.G.G.J. 7788 (BO); Gunung Kelatakan, 2 Aug 1918, Maier Sarip, R. 162
(BO). Java: s.l., 8 May 1931, Clason, E.W. C99 (L); Zollinger, H. 2002 (S). Jawa Timur:
Surabaya, Mantup, Coster, C. s.n. (BO); Gunung Wilis, 1914, Backer, C.A.B. 11469 (BO);
Besuki, Gunung Ringit, Kampong Agoeng, 8 Mar 1940, Buwalda, P. 7542 (BO); Besuki, Jang
Plateau, Djeloewang Ravine, 18 Jul 1938-19 Jul 1938, van Steenis, C.G.G.J. 11079 (BO);
Besuki, Puger, 28 Feb 1940, Buwalda, P. 7225 (BO, L); Nusa Baron, 21 Feb 1845, Zollinger,
H. 2612 (BO); Pasuman, 1 8 Mar 1929, Backer, C.A.B. 36621 (L); Ponorogo to Soemoroto, Jan
1918, Beiimee, J.G.B. 1328 (BO); Soekapoera, Gorge of Oemboelan, 12 Apr 1925, Jeswiet,
J. 670 (WAG). Jawa Tengah: Gunung Lawoe, 1913, Backer, C.A.B. 6607 (L); ibidem, 1913,
Backer, C.A.B. 6687 (BO); Yogyakarta, Gunung Kidoel, 1912, Backer, C.A.B. 2789 (BO, L);
Yogyakarta, Unknown (ex Herb. Lugd. Batav.) s.n. (L); Yogyakarta, Prambanan, Feb 1912,
Ridley, H.N. s.n. (K). Jawa Barat: Pameungpeuk, Goenoeng Karikil, Jan 1933, Jacobson, E.
199 (BO). Nusa Tenggara Barat: Lombok, Sambelia, Gunung Rinjani, 17 Feb 1998, de Wilde,

Revision of Epithema

187

W.JJ.O. & de Wilde-Duyjjes, B.E.E. 21940 (E, L); ibidem, 2 May \96\, Kosterincms, AJ.G.H.
18629 (K, L); Western Sumbawa, Batudulang, Mt Batulante, 10 Apr 1961, Kostermans,
A.J.G.H. 18053B (K); Western Sumbawa, Olat seli, Pemek, 17 May 1961, Kuswata 213 (BO);
Western Sumbawa, Sumbawa Besar, 29 Mar 2004, Hoover, W.S, etal. Dedenl 51 (US); Western
Sumbawa, Sumbawa Besar, Semongkat Atas, 26 Mai* 2004, Hoover, W.S. et al. HW11417 (BO,
US); ibidem, 1 May 1961, Kuswata 105 (BO, L); ibidem, 2 May 1961, Kuswata 121 (K, L).
Nusa Tenggara Timur: Flores, Loeters, JJ. 1625 (L); Flores, Lempe to Waewako, 26 Mar
1974, Schmutz, E. 3623 (L); Flores, Waewako, 21 Feb 1978, Schmutz, E. 4027 (L); Western
Flores, Manggarai, 18 Feb 1979, Schmutz, E. 4345 (L); ibidem, 6 Feb 1981, Schmutz, E. 4732
(L); Timor, Naukae, 29 Jan 1971, Kooy, C.W. 789 (L); Timor, Nikiniki, 20 Feb 1966, Kooy
C.W. 403 (L). Sulawesi Selatan: Bulukumba, Malino, 2 Apr 1922, Bunnemeijer, H.A.B. 10713
(BO, L); South ibidem, 8 Apr 1921, Bunnemeijer, H.A.B. 10890 (BO); Bulukumba, Malino,
Bulutana, 10 Feb 2000, Mendum, M. et al. 0034 (E, SING); Maros, 23 Jan 1998, de Wilde,
W.J.J.O. & de Wilde-Duyfjes, B.E.E. 21918 (E, L).

Notes. Hilliard & Burtt( 1997) descrihedthQ variety Epithema horsfieldiiYar. epiphyllum
which they distinguished from the type variety by the position of the inflorescences. In
Epithema horsfieldii var. horsfieldii they usually arise from the base of the leaf blade
and in E. horsfieldii var. epiphyllum tfoin the midrib, up to 2.3 cm from the base of
the blade. Occasionally the inflorescences of the plants placed in Epithema horsfieldii
var. horsfieldii also arise from the lower midrib but these specimens are more densely
villous inside the corolla than the material placed in E. horsfieldii var. epiphyllum. In
addition the material assigned to Epithema horsfieldii var. horsfieldii is generally more
pubescent than E. horsfieldii var. epiphyllum and has larger and more cucullate bracts.
However, specimens with the morphology of the type variety were found disjunctly in
Java and Timor and that of Epithema horsfieldii var. epiphyllum in Lombok, Sumbawa
and Flores. This would be biogeographically unlikely. The material tfom Sulawesi
is rather different again as much of the material has inflorescences arising in a slight
groove at the base of the leaf blade which is not found in the two existing varieties
whereas other specimens are more similar to the type variety. What is clear is that the
patterns of variation require considerably more study and that if varieties are to be
reeognised they should only be so after more collecting from tliroughout the range of
the species (but particularly in Timor) and after a more thorough understanding of the
variation.

6. Epithema involucratum (Roxb.) B.L.Burtt, Notes Roy. Bot. Gard. Edinburgh 22:
308 (1958). - Gratiola involucrata Roxb., [Hort. Bengal. 80 (1810), nom. nud.] FI.
Indica ed. 1, 138 (1820). - Epithema roxburghii A.DC., Prod. 9: 279. (1845), nom.
superfl. - TYPE: “Moluccas”, Roxburgh s.n. (lectotype PDA, designated here). (Fig.
7)

Aikinia brunonis Wall., PI. Asiat. Rar. 3: 66 (1832); G.Don, Gen. Hist. 4: 665 (1838). -
Epithema brunonis (Wall.) Decne., Nouv. Ann. Mus. Hist. Nat. 3: 404 (1834); Decne.,
Herbarii Timorensis Descriptio 76 (1835); A.DC., Prodr. 9: 279 (1845); Clarke in

188

Card. Bull. Singapore 67(1) 2015

Fig. 7. Distribution of Epithema involucratum (Roxb.) B.L.Burtt (■), Epithema longipetiolatum
(Merr.) Hilliard & B.L.Burtt (A), Epithema longitubum Hilliard & B.L.Burtt (•).

A.DC. & C.DC., Monogr. Phan. 5(1): 179 (1883). - TYPE: Indonesia, East Nusa
Tenggara, Timor, Coepang, Apr 1803, Brown, R. s.n. (lectotype BM, designated here).

Epithema brunonis var. violaceum C.B. Clarke in A.DC. & C.DC., Monogr. Phan. 5(1):
179 {ISS3). - Epithema violaceum Span., Linnaea 15: 330 (1841), nom. mid. - TYPE:
Timor Leste, Spanoghe 58 (holotype P; isotype L).

Herb 4.5-20 cm high, caulescent, indumentum strigose to hispid, hairs sparse to
medium density, to 0.3 mm long on the stem, petioles, peduncles, lower bract, and
the outside of the calyx, to 0.8 mm long on the upper and lower surface of the leaves,
indumentum of lower bract, pedicels and the outside of the calyx also of minute hairs;
stem 1-3 mm wide with 2 to 3 nodes, intemodes 1 .3-3 cm long. Leaves membranous
and petiolate, upper leaves opposite; petioles 0.2^. 5 cm long; blades 1.3-6 x 1-5.4
cm, ovate or cordate, rarely almost oblong symmetrical or barely asymmetrical, apex
rounded to truncate, more rarely broadly acute, base cordate or truncate, inserted evenly
on petiole or not, margin entire or with few, minute teeth, undulate. Inflorescences up
to 5 per plant, tenninal, in lower leaf axils or on base of petioles; peduncles 1 .3-5.5
cm long; bracts cucullate but not always enclosing entire inflorescence, 7-17 x 4-10
mm, margin variable, almost entire to dentate, upper surface sub-glabrous, hairs to 0.2
mm long, placed primarily towards margins of the bract; pedicels 0.9-3. 5 mm long,
indumentum may include strigose hairs to 0.5 mm long. Calyx 2. 9^.4 mm long, tube
1.7-2. 5 X 1-2.5 mm, lobes 1-2.1 x 0.6-1. 9 mm; glabrous inside. Corolla light blue
or bluish to dark violet, tube may be white, 9-11 miu long; tube 5-6 x 1.5-2 mm;
lobes 4-5 X 3-A mm, almost as long as the tube; glabrous outside and inside. Stamen

Revision of Epithema

189

filaments 1 .2 mm long; anther dimensions unknown; staminodes 0.7 mm long. Nectary
of two discrete lobes, 0.6 x 0.6 mm. Ovary cylindrical or sub-cylindrical, 0.8-1 x 0.7
mm, pubescent, hairs dense, hooked, white, to 0. 1 mm long on upper portion of ovary;
style 3.5 mm long, glabrous or sub-glabrous, hairs, when present, hooked and on lower
style; stigma to 0.5 mm wide. Fruit sub-cylindrical to sub-spherical, 1.7-1. 9 x 1.7
mm; operculum 0 . 6 - 0. 8 mm long, sparsely to densely pubescent, hairs hooked, to 0.1
mm long, on entire operculum. Seed fusiform to broadly oval, 0.3-0. 5 x 0. 1-0.2 mm;
pattern straight to spiralled and even.

Distribution. Indonesia (Sulawesi, Timor, Maluku), Timor Leste.

Habitat and ecology. In shaded areas. Highly likely to be lithophytic on limestone and
found in humid places and/or near water.

Provisional lUCN conservation assessment. Data Deficient (DD). The known
localities of this species suggest a fairly widespread distribution. However, it has not
been collected often, many of the collections are fairly old and the exact localities of
some of these older specimens are not known. We, therefore, assign it the status of
Data Deficient until more infomiation becomes available.

Additional specimens examined. INDONESIA: Maluku: Babar Islands, Pulau Weten, 28 Feb
1956, van Borssum Waalkes, J. 3025 (BO, L). Nusa Tenggara Timur: Sumba, Laora, 10 Nov
1925, Iboet 221 (BO); Timor, Spanoghe, J.B. 226 (L); ibidem, Unknown (probably Spanoghe)
226 (L). Sulawesi Tenggarah: Pulau Butung, Baoe Baoe, 20 Feb 1929, Kjellberg, G.K. 314
(BO, S); ibidem, 20 Feb 1929, Kjellberg, G.K. 315 (S).

Notes. key macroscopic features distinguishing Epithema involucratum from
other species are the cordate to ovate leaves with clearly rounded leaf tips and that
it is one of the few species that is glabrous inside the corolla. Epithema longitubum
and some specimens of E. ceylanicum are similar to E. involucratum. The Epithema
ceylanicum specimens can be distinguished from E. involucratum by the presence
of an indumentum inside the corolla, smaller inflorescences and shorter corolla
length. Epithema longitubum can be distinguished by the often longer corolla length
and presence of an indumentum in the corolla. While there were not many corollas
available to examine, it would seem that the lobes of E. involucratum are quite large
in comparison to other species.

Van Borssum Waalkes 3025 (L) from Maluku is rather intennediate between
Epithema involucratum and E. longitubum. It is included in Epithema involucratum
on the basis of its leaf shape, absence of hair in the corolla and the lack of an upper,
solitary leaf It does have medium to high density, strigose hairs on the inner side of the
calyx lobes which is unusual for either species. Its inclusion in Epithema involucratum
significantly extends the known distribution of E. involucratum, as it would also have
done for E. longitubum if it were included there.

Kjellberg 3 14 (S) from Sulawesi also extends the range of Epithema involucratum
although, while most of its characters are typical for E. involucratum, it has hooked
hairs at medium density on the ovary which is unusual for this species.

190

Card. Bull. Singapore 67(1) 2015

Epithema brunonis var. violaceum is included in synonymy of E. involucratum.
Clarke ( 1 883) used depth of flower colour, length of the corolla and the leaf pubescence
to delineate it from the type variety. The flower colour and leaf pubescence is not
different from that seen in other E. involucratum specimens and the corolla length,
from material that Clarke himself examined, is 9 mm not 12 mm as Clarke stated when
differentiating the variety. The key differences are nominal; the shape of the base of
a couple of leaves differ slightly and the peduncles can be up to 15 mm longer than
in typical E. involucratum. Epithema brunonis var. violaceum, however, is within the
range of variation for^. involucratum.

In the protologue of Aikinia brunonis there is no indication of a collector of
the specimen used to make the description, only that it is based on a collection from
Coepang (now called Kupang) in Timor which was flowering and fruiting in April
1803. There is a Robert Brown collection from Coepang collected in April 1803 in the
BM. This specimen is lectotypified here.

7. Epithema longipetiolatum (Merr.) Hilliard & B.L.Burtt, Edinburgh J. Bot. 54: 112
(1997). - Epithema brunonis var. longipetiolatum Mem, Philipp. Joum. Sci., C. 11:
313 (191 6). - TYPE: Indonesia, Maluku, Ambon, Halong, 50-100 m, 26 September
1913, Robinson, C.B. 1727 (lectotype L, designated here; isolectotypes BO, K, US).
(Fig. 7)

Herb 5-30 cm high, acaulescent or caulescent; stem 2-3 mm wide with two nodes,
intemodes 7-10 cm long. Leaves membranous, petiolate, usually solitaiy but upper
leaves may be opposite, opposite leaves are occasionally unequal in size; petioles (0-
)0.5-12(-l 8) cm; blade 3-13.6 x 2.4-10.2 cm, cordate to broadly ovate or occasionally
sub-lanceolate, symmetrical or asymmetrical, apex acute, base sub-cordate or sub-
auriculate to cuneate, inserted evenly on petiole or distinctly offset, margin almost
entire to semulate or denticulate, veins often widely spaced; upper surface glabrous or
sparsely to densely strigose, hairs to 0.4 mm long, with or without a minute hooked
pubescence; lower surface finely pubescent, at medium to high density, surface often
appearing fuzzy. Inflorescences 1-6 per plant; peduncles 0.2-6 cm long, terminal or
in the axils of upper leaves; bracts cucullate but not enclosing the entire inflorescence,
5-12 X 4-9 mm, margin somewhat dentate to dentate and serrate, upper surface
sub-glabrous to sparsely strigose, hairs straight, to 0.2 mm long, denser towards the
margin; pedicels 1-4.2 nun long. Calyx 2. 7-5. 6 mm long; tube 1.5-5 x 1.5-3. 8 mm,
lobes 1.2-2. 7 x 0.6-1. 7 mm; minutely pubescent (often finely) outside, occasionally
strigose, at medium to high density, all hairs hooked or primarily hooked and to 0.25
mm long, hairs on inside of lobe tips to 0.13 mm, occasionally sub-glabrous. Corolla
tube white, lobes lilac or pale lilac, lobes with darker coloured but variable markings,
5.5-10.2 mm long; tube cylindrical, 4.2-6.4 x 1.5-1. 9 rmu, lobes 1.2-3. 8 rmu long;
with a band of villous hairs inside around middle, up to 1 mm wide. Stamens 2. 1-3.1
mm long; filaments 1.5-2. 3 mm long; anthers 0.6-0. 8 mm long; staminodes 1.2-2. 3
mm long. Nectary of two discrete lobes, margin entire or undulate, 0. 5-1.1 x 0.7-1. 7

Revision of Epithema

191

mm. Ovary sub-cylindrical or sub-spherical, 1-1.5 x 0.7-1. 3 mm, densely pilose,
hairs usually straight or occasionally straight and hooked, placed on upper portion of
ovai*y, 0.04-0.08(-0.25) mm long; style 3. 1^,4 mm long, wider at base to 0.33 nuu,
style glabrous or with very few hairs at base, these straight, to 0.17 mm long; stigma
0.4 mm wide. Fruit obovate to sub-cylindrical, 2-A x 2-3.5 mm; operculum 0.6-1 .9
mm, finely setose or pilose, hairs medium to high density, as on ovary. Seed narrowly
to broadly ovoid, 0.3-0. 5 x 0. 1-0.2 mm, light to medium brown, pattern straight to
partially spiralled, often with thick, clearly defined walls.

Distribution. Indonesia (Ambon, Bum, Seram, Am, Sulawesi, Papua).

Habitat and ecology. Lithophytic on limestone. In humid, shaded places, often on
rocks in stream beds at 0-1770 m altitude. Flowering and fmiting throughout the year.

Provisional lUCN conseiyation assessment. Least Concern (LC). This species
is widespread. However, as it fi'equently occurs on karst limestone, which is often
conuuercially exploited, its status should be monitored.

Additional specimens examined. INDONESIA; Maluku: Am Islands, Pulau Kobroor,
Namadoeboele, 1 0 Jun 1938, Biiwalda, P. 5202 (BO); Am Islands, Pulau Wokam, Dosinamalaoe,
1 Jun 1938, Buwalda, P. 5110 (A, BO, K, L); Bum, NW Bum, Bara, 5 Dec 1984, van Balgooy,
M.M.J. 5066 (BO, L); Buai, NW Bum, Bara, Waeduna River, Nooteboom, H.P. 5217 (L);
ibidem, 25 Nov 1984, van Balgooy, M.M.J. 4889 (E, L); Ambon, Boerlage, J.G. 528 (BO);
ibidem, Teijsmann, J.E. s.n. (BO); Seram, 9 Jan 1938, Eyma, P.J. 25 74 (BO, L); Seram,
Kampong Kwaos, Gunung Toenlean, 8 Sep 1938, Buwalda, P. 6006 (K, L); Seram, Kampong
Selagor, Gunung Selagor, 26 Aug 1938, Buwalda, P 5748 (BO, L); Seram, Kecamatan Seram
Utara, Goa Pohon Damar to Gunung Roihelu, 24 Jan 1985, Kato, M. et al. C.5232 (L); Seram,
Kecamatan Seram Utara, Goa Pohon Damar to Sawai, 27 Jan 1985, Kato, M. et al. C.5728
(L); Seram, Pileana, 28 Oct 1937, Eyma, PJ. 1826 (A, BO, K, L, SING); Seram, 4 Feb 1918,
Kornassi 967 (BO, L); Seram, Manusela National Park, Gunung Binaia, 26 Aug 1987, Argent,
G.C.G. C8754a (E); Seram, Tehoru, Hatumete Pass, 29 Nov 1917, Kornassi 621 (BO, L);
Seram, Tehom, Saunulu, Murkele Ridge, 21 Jul 1986, Kato, M. etal. C, 11 441 (L); ibidem, 21
Feb 1985, Kato, M. et al. C.5144 (L); Seram, Taniwel, Between Buria and Wae Mala, 3 Feb
1985, Kato, M. et al. C.6007 (A, L). Papua Barat: Kabupaten Raja Ampat, Waigeo Island,
Tuluk Mayalipit, Warsamdin, 1 May 2008, Okada, H. & Tsukaya, H. OT-48 (BO); Biak, Aet
& Idjan 942 (BO, L); Biak, Gunung Wawah, 22 Jul 1939, Aet & Idjan 32 (A, BO, K, L); Fak-
Fak, Fak-Fak River, 23 Feb 1962, Vink, W. BW12141 (L). Sulawesi Tengah: Soroako, Mt.
Wawonsem, 2 Jul 1 979, Hennipman, E. 610g (A, E, L).

Notes. The specimens of Epithema longipetiolatum with leaves arising from ground
level could be confused with E. rennellense and E. strigosum although there is no
overlap in distribution. Leaves arising from ground level are always present in E.
rennellense and E. strigosum and opposite leaves are never seen in these species. The
leaf margin of Epithema strigosum has large wide teeth (entire to weakly dentate in E.
longipetiolatum). Epithema rennellense has straight and hooked hairs on the ovary and
operculum (usually only straight in E. longipetiolatum).

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‘Loiigipetiolatiim’ refers as much to the length of the secondary intemode as
to the petioles, giving the plant an elongate appearance. For example, in Kornassi
967 (L), the secondai*y internodes are a third of the total plant height and equal to the
primary intemode and the upper petiole and blade combined.

Epithema longipetiolatum may have solitary leaves only, opposite leaves only,
or opposite and solitaiy leaves. This does not coirelate with geography and both types
can be found within a population. For example, duplicates of the type collection have
only opposite leaves or only solitary leaves while Argent C8754a (E) from Seram
includes plants that have only opposite leaves or both opposite and solitary leaves.

Some specimens of Epithema longipetiolatum are similar to specimens of E.
henthamii. For example, one duplicate of Eyma 1826 (L), from Seram, has a similar
leaf shape and margin to E. benthamU. We have seen no specimens of Epithema
benthamii from Seram, but E. longipetiolatum and E. benthamU are found in the same
locality in the Waeduna River district on Burn Island. The ovary and operculum hair
of specimens of Epithema longipetiolatum found on Burn Island often have a mixture
of long (to 0.25 mm), almost villous hairs that are straight or straight and hooked like
E. benthamii, in addition to the expected, short hairs. This contrasts with the type
specimen of Epithema longipetiolatum from Ambon, theF^. longipetiolatum specimens
from Seram, Wokam Island and the Aet & Idjan 942 (L) specimen from Biak in West
Papua, which have minute, pilose, straight hairs (rarely with a few hooked hairs) on
the ovary and operculum. It is possible some hybridisation occurs between Epithema
longipetiolatum and E. benthamii in these localities.

The minute hooked pubescence on the pedicels and outer calyx is quite
distinctive, especially for the specimens from Seram. For most specimens of Epithema
longipetiolatum, the indumentum of the upper leaf surface is of long, strigose hairs and
looks somewhat like the indumentum of the upper surface of the leaves of E. benthamii.
Specimens from Bum, however, have a dense, uniform and short indumentum, giving
the leaves a fuzzy look.

8. Epithema longitubum Hilliard & B.L.Burtt, Edinburgh J. Bot. 54: 112 (1997). -
TYPE: Timor, Nasinutan, 11 March 1939, Bloembergen, S. 3458 (holotype L; isotype
BO (image seen)). (Fig. 7)

Herb 7.5-20 cm high, with a distinct (but sometimes short) stem before first leaf; stem
0.6-2. 5 mm wide with two nodes, intemodes 1.5-7 cm long. Leaves membranous,
petiolate, upper leaves opposite or with an uppermost leaf solitary opposite a peduncle;
petiole of lowest leaf 4-6 cm long, petioles of upper leaves 0.5-5. 5 cm; blade 1 .7-11
X 1-7.8 cm, sub-orbicular to broadly ovate or somewhat oblong, symmetrical or not, if
not one side up to 1 .3 times wider than the other, apex rounded to acute, base shallowly
auriculate or cordate to rounded, inserted evenly on the petiole or not, margin almost
entire to sermlate or denticulate, venation sub-opposite towards base, usually alternate
towards leaf tip, widely spaced; upper surface with a longer indumentum of hairs to
1 mm long, with or without an additional shorter pubescence of hooked hairs; lower

Revision of Epithema

193

surface drying pale, indumentum as upper surface but less dense. Inflorescences 2-10
per plant; peduncles 0.5-8 cm long, tenninal or in leaf axils; bracts diamond-shaped
or trullate, cucullate but only enclosing a small portion of the inflorescence, 2-9 x
3-6 nmi, margin roughly entire with occasional dentation or with some crenate and
dentate lobes, indumentum of lower surface variable with hairs to 0.5 mm long; upper
surface sub-glabrous, sparsely and minutely pubescent or sparsely strigose; pedicels
0. 75^.2 mm long. Calyx 2.9-5.2(-7) mm long; tube 1. 4-4.2 x 1.3-2. 5 mm; lobes
1-2.9 X 0.4-1 .3 mm; minutely and sparsely to densely pubescent outside and with
or without larger hairs to 0.7 mm long at medium to high density throughout calyx
or largely on lobes; glabrous to sparsely and minutely pubescent inside on upper part
of lobes. Corolla blue-purple, c. 10-17 mm long; tube cylindrical, 8-12 x 1.3-2 mm,
up to 7 times as long as wide; lobes 2-5 mm long; with band of villous hair inside
in lower part. Stamens 1.8-2. 3 mm long; filaments 1-1.5 mm long; anthers 0.8 mm
long; staminodes 0.6-1 .4 mm long. Nectaiy of one or two lobes, partly to entirely
encircling ovary, 0. 5-0.9 x 1.3-1 .5 mm, margin undulate and of varying height. Ovary
sub-cylindrical or broadly ovoid with a truncate base, 0. 9-1.1 x 0.4— 0.6 mm, densely
pubescent on upper portion, hairs hooked; style 8.8-10 mm long, glabrous, rarely
with few hooked hairs at base. Fruit sub-spherical to cylindrical, 1.3-2. 5 x 1.7-2
mm; operculum 0. 5-0.6 mm, pubescent tliroughout but denser at top, hairs hooked, to
0.08 mm long. Seed ovoid to broadly ovoid, 0.3-0. 5 x 0. 1-0.2 mm, pattern partially
spiralled to spiralled, regular.

Distribution. Indonesia (Timor, Flores).

Habitat and ecology. Lithophytic on limestone in humid, shady places near or beside
streams and cave entrances at 900-1200 m altitude. Flowering and fruiting March and
May.

Provisional lUCN consen’ation assessment. Data Deficient (DD). The most recent
collection of this species is from 1974 and the collection density on both Flores and
Timor is too low to have an accurate idea of the curtent distribution.

Additional specimens examined. INDONESIA: Nusa Tenggara Timur: Flores, 25 Mar 1972,
Verheijen, J.A.J. 3070 (L); ibidem, Mar 1974, Verheijen, J.A.J. 3323 (L); Flores, Ruteng-Mano,
16 May 1973, Schmutz, E.3208 (L); Flores, Ruteng-Reo, 7 Mar 1968, Schmiitz, E.2127 (L);
Timor, 30 Jan 1929, Walsh, M.E. 62 (BO); Timor, Nasinutan, 17 Mar 1939, Bloembergen, S.
3496 (BO, L).

Notes. This species is most similar to Epithema involucratum but the inflorescences,
bract and corollas are usually much larger. Unfortunately the locality information on
specimens of both Epithema longitubum and E. involucratum is poor and, therefore, it
is as yet unclear whether these two species co-occur or not without further field work.

Although nectary characters have not proven useful in Epithema, and relatively
few specimens have been examined for this species, it is interesting to note that the

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Card. Bull. Singapore 67(1) 2015

specimens from Flores have two nectary lobes and the specimen from Timor has one.
The sampling density, however, is currently too low to make any firm conclusions.

9. Epithema madulidii Hilliard & B.L.Burtt, Edinburgh J. Bot. 54: 112 (1997). -
TYPE: Philippines, Coron Island, Banol, 30 Sep 1993, Madulid, D.A., Agoo, Reynoso
& Puentes, R. 11563 (holotype PNH; isotype E). (Fig. 6)

Herb 13.2-50 cm high, caulescent; stem 1^.2 mm wide with two nodes, intemodes
1.5-11 cm long. Leaves membranous or strongly membranous, lower leaf petiolate,
upper leaves sessile; lower petioles 1-10 cm long, sparsely to densely hairy, often
with two distinct lengths of hair, hairs white, to 1.1 mm long; lower leaf blade 9-13 x
4.2-7.2 cm, upper leaves 2-4.5 x 1.7-3. 5 cm, lower leaf cordate to ovate, base cordate
to rounded, upper leaves ovate to elliptic, all leaves usually asymmetrical, one side
1. 1-1.8 times wider than the other, apex broadly acute to rarely rounded, base cordate
to obtuse, margin sub-entire to sinuate to minutely dentate, inserted evenly on petiole
or not; upper surface sparsely to densely hairy, often with two distinct lengths of hair,
hairs to 0.4 mm long; lower surface similar, hairs to 0.5 mm long, indumentum on veins
often white and to 0.8 mm long. Inflorescences 1-3 per plant, tenninal; peduncles
1.4-8 cm long; bracts whitish at base, upper half violet, cucullate, partially enclosing
inflorescence, 10-25 x 6-15 mm, margin dentate to irregularly dentate, lower surface
sub-glabrous or sparsely strigose, upper surface glabrous; pedicels 1^.2 mm long.
Calyx 6.3-7 imn long; tube 3-5.2 x 2. 1-3.8 nmi, often drying a lighter colour than the
lobes; lobes 1.5-2. 9 x 1,1-3. 8 mm, outside glabrous or with sparse hairs primarily on
lobes, hairs to 0.6 mm long; sub-glabrous or pubescent inside, hairs straight, to 0.1 mm
long, mostly only in lobes, occasionally in the tube. Corolla violet at tips, 5-10.4 mm
long; tube cylindrical, 3. 3-6. 5 x 1 .5-1 .8 mm; lobes 1 .7-4.5 mm long; sub-glabrous to
pubescent on lobes outside, hairs hooked or straight, glabrous inside. Stamens 1.3-1. 5
mm long; filaments 0.6-0. 7 mm long; anthers 0.7-0. 8 mm long; staminodes 0.6-0.75
mm. Nectary of one lobe, encircling half to two-thirds of the ovary, 0.5-0. 9 mm high,
margin entire or almost entire. Ovary sub-spherical, 0.7-1. 1 x 0.5-0. 8 mm, densely
pubescent on upper portion of ovaiy, hairs hooked, white, to 0.04 mm long; style 6.3
mm long, glabrous or occasionally with few straight hairs. Fruit obovate to obovate-
cylindrical, 2.6^ x 2-2.7 imu; operculum 0.6-0. 9 mm, sub-glabrous to densely
pubescent, hairs straight and hooked, to 0.13 mm long. Seed ovoid or with one side
flattened, 0. 5-0.6 x 0. 1-0.2 mm, medium to dark brown, pattern more or less straight
to spiralled, regular, lines of pattern thickened.

Distribution. Philippines (Coron Island).

Habitat and ecology. Found in shallow soil in limestone rock crevices in humid,
shaded areas at c. 50-550 m.

Provisional lUCN conservation assessment. Data Deficient (DD). This species is only
known from two eollections from Coron Island and has both an EOO and AOO that

Revision of Epithema

195

would put it in the Critically Endangered category. However, this limestone island and
neighbouring islands are not well collected and the impact of tourism on the habitat is
not clear.

Additional specimen examined. PHILIPPINES: Palawan: Coron Island, Sep 1922, Lopez, G.
BS41343 (BO, K, L, P, US).

Notes. This species is known from two collections from Coron Island collected in 1922
and 1993. Due to the poor condition of the specimens, particularly the Lopez BS41343
collection, some information such as leaf blade, base and tip shape is assessed from
very few leaves.

The calyx of this species is quite large and the tube dries a very light brown
colour in comparison to the lobes although this is based on a very small sample. The
corolla tube is glabrous, or appears so, and, in combination with the light colour and
large size of the calyx, is useful for identification from dried specimens. It is similar to
the sessile-leaved form of Epithema saxatile but can be distinguished by the complete
lack of hair inside the corolla and the presence of straight hairs on the operculum. In
addition the calyx is larger (6.3-7 mm versus 3.3-5 mm) and there are hairs on the
inside of the calyx lobes (lacking in Epithema saxatile).

10. Epithema membranaceum (King) Kiew, Malayan Naturalist J. 38(3): 36 (1985);
B.L.Burtt, Thai For. Bull. (Bot.) 29: 93 (2001). - Argostemma membranaceum King,
J. Asiat. Soc. Bengal, Pt. 2, 72: 145 (1903); Ridley, FI. Malay Penins. 2: 24 (1923),
- TYPE: Malaysia, Penang, Muka Head, Curtis 955 (lectotype SING, designated by
Kiew (1985)). (Fig. 8, 9)

Herb 10-30 cm high, caulescent, indumentum of minute hooked pubescence and
strigose hairs, usually sub-glabrous or sparse; stem 1.5 mm wide, can be red-brown in
colour, with 2-3 nodes, intemodes 0.6-12 cm long. Leaves often finely membranous,
petiolate, upper leaves opposite, infrequently upper leaves solitary or sub-opposite;
petioles 0.8-14.5 cm long; blade 3.6-11.5 x 3-10.5 cm, broadly ovate or cordate,
occasionally sub-orbicular or almost oblong, symmetrical or barely asymmetrical,
apex acute to rounded, upper portion of leaf sometimes curved, base sub-auriculate
to truncate, inserted evenly on petiole, margin entire to seiTate, crenate or with large,
rounded dentation; upper surface glabrous to sparsely strigose (occasionally to medium
density), hairs to 0.6 mm long; lower surface glabrous, sub-glabrous or covered
in dense, minute hooked pubescence, giving a fuzzy appearance. Inflorescences
(l-)3-9 per plant; peduncles 0.5-17 cm long, primarily from terminal leaf axils,
rarely from the base of the petioles; bracts usually small, obovate, ovate or elliptic,
not enclosing inflorescence, if sub-cucullate then only enclosing a small proportion
of the inflorescence, 2-11 x 3-12 mm, margm entire or with limited dentation,
lower surface sub-glabrous, upper surface glabrous or with sparse hair towards bract
margin; pedicels 1-3.5 mm long, densely and minutely pubescent, with or without
few additional straight, strigose hairs to 0.3 mm long. Calyx 2. 1-5.2 mm long; tube

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Card. Bull. Singapore 67(1) 2015

Fig. 8. Epithema membranaceum (King) Kiew A. Habit. B. Inflorescence. C. Calyx. D.
Calyx indumentum detail. E. Flower opened out. F. Stigma lateral view. G. Fruit showing
seeds, placenta and operculum. H. Operculum indumentum. I. Seeds. Scale bars: A = 4
cm, B = 5 mm, C = 3 mm, D, H, I = 0.5 mm, E = 4 mm, F = 1 mm, G = 2 mm. Drawn by
Claire Banks from Larsen et al 42245 (A-D, G-I) and Larsen et al. 42312 (E, F).

Revision of Epithema

197

Fig. 9. Distribution of Epithema membranaceum (King) Kiew (•) Epithema sarawakense
Hilliard & B.L.Burtt ( A).

1.25-3.5 X 1.3-2. 3 mm; lobes 0.6-3. 1 x 0.75-2.5 mm; indumentum outside as on
pedicel, sub-glabrous inside, a few hairs in lobes. Corolla colour variable, white, pale
pink, pale violet, pale blue or purple or whitish-purple with purple markings, 3.5-7
mm long; tube 2.5-5 x 2 mm, lobes 1-1.9 mm long; pubescent outside, hairs hooked
and straight to 0.1 mm long, with a band of villous hair inside. Stamen filaments
0.75-0.8 mm long; anther characters unknown; staminodes 0.6 mm long. Nectaiy of
one lobe almost or entirely encircling ovary, 0.75-1 x 2.2 nun. Ovary spherical or sub-
spherical, 0.75-1 X 0.6-1 mm, pubescent, hairs medium to high density, fine, hooked,
rarely with one or two straight hairs, hyaline, on upper portion of ovary; style 2-3.8
mm long, occasionally glabrous, pubescent on lower to mid-style, hairs hooked, to
0.13 imn long; stigma c. 0.4 mm wide. Fruit obovate to cylindrical, 2-3 x 2-3 mm;
operculum 1 mm long, pubescence as for ovary. Seeds narrowly ovoid to ovoid, 0.3-
0.5 X 0. 1-0.2 mm, dark brown, pattern straight to spiralled, regular.

Distribution. Thailand, Peninsular Malaysia, Indonesia (Sumatra).

Habitat and ecology. Primarily on limestone, occasionally granite, in shaded areas of
evergreen forests near waterfalls or streams. Found in lowland dipterocarp forests in
Malaysia at 30-415 m altitude. Flowering and fruiting Febmary, April-December.

Provisional lUCN consefyation assessment. Least Concern (LC). This species is
common and widespread although, as it mostly occurs on limestone and limestone
habitats which are often threatened, its status should be monitored.

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Card. Bull. Singapore 67(1) 2015

Additional specimens examined. THAILAND: Chachoeiigsao: Kliao Tak Groep, 6 Nov 1993,
Larsen, K. etal. 44271 (AAU), Nakhon Si Thammarat: Klia Noni, Khuan Thong, Khao Krod
Cave, 20 Dec 2006, Pooma, R. et al. 6486 (E); Nop Phitum, Khao Luang National Park, Khao
Luang, 25 Oct 1991, Larsen, K. et al. 42584 (AALf); ibidem, 25 Oct 1991, Larsen, K. et al.
42579 (AAU). Narathiwat: 21 Dec 1968, Phusomsaeng, S. etal. 19 (BKF, E); Bacho National
Park, 11 Oct 1991, Larsen, K. et al. 42212 (AAU, P); Sungai Padi, Chatvarin Falls, 18 Oct
1970-19 Oct 1970, Charoenphol, C. et al 3976 (AAU, E, K, L); Pattani, Kliao Chai Son, 23
Dec 2006, Pooma, R. et al 6578 (E); Pattani, Sai Khao Waterfall, 9 Oct 1991, Larsen, K. et
al. 42245 (AAU); ibidem, 20 Dec 1972, Santisuk, T. & B.N. 451 (BKF). Phatthalung: Khao
Chai Son, Road to Than Nam Yen, 23 Dec 2006, Pooma, R. et al. 6579 (BKF); Kao Ok Thalu,
21 Apr 1928, Kerr, A.F.G. 15775 (K). Songkhla: Kliao Chang Lon, 24 Jul 1928, Kerr, A.F.G.
15893 (K); Saba Yoi, Thani Ru Nok Sak (Cave), 21 Oct 1991 , Larsen, K. et al. 42471 (AAU,
BKF); Tam ta Lord, 25 Nov 1990, Larsen, K. et al. 41713 (AAU). Surat Thani: Khan Thuli,
6 Sep 1931, Put 4118 (K). Yala: Bannang Sata, 26 Nov 1961, Suvanakoses, P. 1727 (BKF, K,

L, P); Nam Tok Than To Falls, 10 Dec 1966, Sangkhacliand, B. 1405 (BKF); Tahndo, Tahndo
Falls Forest Park, 1 2 Nov 1 986, Maxw>ell, J.F 86-893 (L, PSU); Than To, Bang Lang National
Park, 17 Jul 2004, Pooma, R. et al. 4298 (BKF).

MALAYSIA: Peninsular Malaysia: Kedah: Baling, 24 Apr 1987, Stone, B.C. et al. 870424-
5/1 (KLU); Kelantan: Gua Musang, 11 Aug 1971, Chin, S.C. 1404 (KLU); ibidem, 1 Nov
1 967, Shimizu, T. & Stone, B.C. 74427 (KYO); Kuala Beds, 15 Feb 2003, Kiew, R. RK5254 (E,
SING). Pahang: Bentung, Bukit Chintamani, 3 Oct 1931., Henderson, M.R. 25010 (BO, SING);
Gua Tipus, 10 Aug 1929, Henderson, M.R. 22580 (SING); ibidem, 15 Oct 1927, Henderson,

M. R. 19406 (BO); Gunung Senyum, 30 Jul 1929, Henderson, M.R. 22382 (BO, SING); ibidem,
28 Nov 1984, Kiew, R. RK1590A (UPM); Kota Gelanggi, 28 Nov 1984, Kiew, R. RK1584
(UPM); Merapoh, Gua Layang, 13 Aug 1971, Chin, S.C. 1514 (E, K); Taman Negara, Batu
Luas, 21 Oct 1982, Kmv, R. RK1231 (UPM); Taman Negara, Batu Luas, Gua Daun, 1 Oct
1984, Kiew, R. RK1347 (UPM); Taman Negara, Gua Luas, Wilkie, P. FRI52899 (E). Penang:
1822, Wallich, N. 9080 (K). Perak: Grik, Temenggor Dam, 9 Sep 1993, Turner, I.M. & Yong
158 (SING); Grik, Temenggor Dam, Sungei Singor, 8 Nov 1993, Saw, L.G. FR139945 (K).
INDONESIA: Sumatera Utara: Asahan, 12 May 1927-21 May 1927, Bartlett, HH. 7740
(US).

Notes. This species is usually quite delicate and distinctive in its vegetative form.
Usually the lower and middle petioles are 0.7-1. 5 times the length of the leaves
which are all cordate-ovate with sub-cordate to sub-auriculate leaf bases. The petiole/
leaf length ratio decreases up the plant as in all petiolate species of Epithema, but
even the uppemiost petioles often appear long in comparison to other species. The
curved asyimnetric upper leaves of some plants are also seen in Epithema steensii and
occasionally in E. tenerum. While peduncles usually originate in the axils of temiinal
leaves, a small number of specimens also have peduncles arising from the base of the
petioles of the terminal leaves.

Epithema membranaceum can be similar to E. parvibracteatum and more rarely
to E. saxatile. The similarity to Epithema parvibracteatum is discussed under that
species. Label data suggests that Epithema membranaceum co-occurs with E. saxatile
on Gunung Senyum and at Batu Luas, both in Pahang, Peninsular Malaysia, Epithema
membranaceum can be separated from E. saxatile by the size of the bract, pubescence
on the style and shorter hairs on the ovary/operculum.

Revision of Epithema

199

11 . Epithema paimbracteatum Hilliard & B.L.Burtt, Edinburgh J. Bot. 54: 112 (1997).
- TYPE: Malaysia, Selangor, Batu Caves, 60 m, 3 November 1953, Sinclair, J. 7821
(holotype E). (Fig. 10)

Herb 6.5-19.6 cm high, caulescent, indumentum of strigose and setose to hispid hairs,
to 0. 8-1.1 mm long on stem, petioles, upper leaf surfaces and peduncles; stem 0.8-
4.4 mm wide with two nodes, internodes 1.2-3. 5 cm long. Leaves membranous or
strongly membranous, petiolate; petiole of lowest leaf 0.8-5 cm long, upper petioles
0.4-2 cm long, densely hairy; blade of the lowest leaf 2.6-1 1.3 x 1.8-8 cm, upper leaf
blades 2. 3-8. 5 x 1.3-6 cm, leaves frequently asymmetrical, one side up to 1.4 times
wider than the other, apex acute to broadly acute, base shallowly auriculate to obtuse,
inserted evenly on petiole, margin sub-entire to dentate, bidentate or serrate; upper
surface often with two clearly different lengths of hair at medium to high density;
lower surface drying white-green, longest hairs primarily on veins and at margins.
Inflorescences 1-5 per plant; peduncles (0.2-)0.5-10.8 cm long, terminal and in leaf
axils, hispid; bracts small, rarely sub-cucullate or trullate, 1-7 x 2-7 mm, margin often
deeply lobed with 3 lobes, but also dentate; lower surface densely hairy with hairs to
0.6 mm long, upper surface with primarily straight hairs to 0.6 mm long, mostly denser
and longer towards margins; pedicels 0. 8-4.2 mm long, sparsely to densely hairy, hairs
hooked and to 0.2 mm long, also with or without longer straight hairs to 0.6 mm long.
Calyx 3. 5-4. 3 mm; tube 1.5-2. 9 x \ .1-2.1 mm, lobes 0.8 -2.9 x 0.6-1. 9 nun; densely
hispid throughout or mainly on lobes outside, hairs to 0.7 mm long; pubescent inside,
hairs straight, on lobes. Corolla white, 4.2-5.4 mm long; tube cylindrical, 2.9-3. 8 x
1-1.5 mm; lobes 0.8-1. 3 mm long, margin entire; with few hooked hairs inside or
with ring of villous hair. Stamens 1. 1-1.6 mm long; filaments 0.6-1 mm long; anthers
0.5-0. 6 mm long; staminodes 0.75-1 mm long. Nectary 1-3 discrete lobes, if tlnee
the third lobe is small, margin almost entire or undulate, 0.6-1. 3 x 0.3-1 .9 mm. Ovary
spherical or sub-cylindrical, 1-1.3 x 0.9-1 mm, sub-glabrous to densely pubescent
or pilose, hairs straight and/or hooked, dense on upper ovary, to 0.2 nuu long; style
1. 9-3.2 mm long, glabrous or with few straight and/or hooked hairs; stigma c. 0.3 mm
wide. Frw/f obovate to cylindrical to sub-spherical, 1,5-2. 8 x 1.9-2, 5 mm; operculum
0.4-0. 9 mm long, indumentum as on ovary, hairs may be densest at the very top of the
ovary. Seed narrowly to broadly ovoid, 0.3-0. 5 x 0. 1-0.2 mm, light brown to medium
brown, pattern almost straight to spiralled, reasonably regular.

Distribution. Peninsular Malaysia.

Habitat and ecology. On limestone or other rocks, in shade, often in crevices of
limestone, at 60-1 50 m altitude. Flowering and fruiting February, May, July-November.

Provisional lUCN conservation assessment. Least Concern (EC). This species is fairly
widespread in Peninsular Malaysia although, as it mostly occurs on limestone and
limestone habitats which are often threatened, its status should be monitored.

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Fig. 10. Distribution of Epithema parvibracteatum Hilliard & B.L.Burtt (•), Epithema steenisii
Hilliard & B.L.Burtt (A) and Epithema strigosum (C.B. Clarke) Hilliard & B.L.Burtt (■).

Additional specimens examined. MALAYSIA: Peninsular Malaysia: Kelantan: UluKelantan,
Bertram, 30 Jul 1962, Unknown 147 (K, L), Pahang: Kota Gelanggi, Aug 1891, Ridley, H.N.
s.n. (SING); Kuantan, Kuantan-Keniaman Road, 19 Jul 1979, Kochummen, K.M. FR126221
(L); Taman Negara, Batu Subuh, 8 Oct 1984, Kmy R. RK1498 (UPM), Perak: Hulu Perak,
Lenggong, Kpg. Gua Badak, 28 Oct 2008, Imin, K. FRI63208 (SING); Kamuning, Feb 1904,
Ridley, H.N. 11883 (K). Selangor: Kanching, Bukit Anak Taloin, 4 Oct 1969, Stone, B.C. 8830
(KLU); ibidem, 29 May 1970, Stone, B.C. & Mahmud 8400 (KLU). Selangor: Kanching,
Bukit Takun, 27 Sep 1970, Chin, S.C. 392 (KLU); ibidem, 24 Oct 1967, Shimizu, T. & Stone,
B.C. 13739 (KYO).

Notes. The distinguishing characters for this species are a small bract and the dense
and long hair covering, particularly on the peduncles, pedicels and calyx. In addition,
the corolla is quite small, only 4.2-5. 4 mm long. There is considerable variation in the
ovary/operculum hair. This variation was observed both within a single inflorescence
and between specimens. Variation in the type of hair found on the inside of the corolla
is unusual and, apart from Epithema parvibracteum, is only seen in E. horsfieldii and
E. rennellense.

The larger plants are rather similar to Epithema membranaceum and some
also to E. saxatile. The specimens of Epithema parvibracteatum, however, have an
extremely reduced bract and a hispid indumentum on the inflorescence.

12. Epithema philippinum (Hilliard & B.L.Burtt) Bransgrove, stat. nov. - Epithema
strigosum philippinum Hilliard & B.L.Burtt, Edinburgh J. Bot. 54: 113 (1997).

Revision of Epithema

201

- TYPE: Philippines, Mindanao, Surigao, Mt Kabatuan, 83 m, 18 March 1949,
Mendoza, D.R. & Convocar, RR 297 [PNH10434] (holotype PNH; isotype A). (Fig. 6)
Herb 6-13 cm high, usually acaulescent with leaves arising near gi'ound level; stem
1-2.5 mm wide with 0-1 nodes, intemode if present 1.4-2. 3 cm long; stem densely
pubescent, hairs to 0.5 mm long. Leaves strongly membranous, petiolate, with one or
more inflorescences opposite each solitary leaf; petioles 2-20 cm long; blade 3-11
X 2.4-4.4 cm, cordate, symmetrical, apex and tip acute, base cordate to truncate,
inserted evenly on petiole, margin dentate and/or bidentate or crenate; upper surface
weakly to strongly strigose, sometimes with setose hairs, hairs straight, hooked or
a combination of both, to 0.7 mm long; lower surface pubescent, hairs to 0.4 mm
long. Inflorescences 1-2 per plant; peduncles 1.6-10.5 cm long, originating opposite
solitary leaves, strigose and setose, haii's dense, to 0.6 mm long; bracts cucullate but
not enclosing entire inflorescence, 3-8 x 3-8 mm, margin dentate; lower surface
strigose, sometimes with two distinct lengths of hair or layers of hair visible, hairs to
0.4 mm long; upper surface strigose, on entire surface but denser towards margins,
hairs to 0.3 mm long; pedicels 1.2-3. 8 mm long, pubescent or hispid, hairs primarily
straight, to 0.4 mm long. Calyx cylindrical, 3. 5-5. 5 mm long; tube 2.3-3 x 1.5-3 mm;
lobes 1.2-2. 5 x 0.8-1. 3 mm; outside with a minute hooked pubescence and longer,
primarily straight hairs covering the entire calyx or only on lobes, straight hairs 0.2-
0.4 mm long, hooked hairs 0.13-0.3 mm long; hairs on lobes inside. Corolla bluish,
5-6 mm long; tube cylindrical, 3. 7-4.2 x 1.2-3 mm; lobes 1.5-2 mm long; band of
villous hair inside in upper tube. Stamens 1-1.5 mm long; filaments c. 0.7-1 mm
long; anthers c. 0.5 mm long; staminodes 0.7-1 mm long. Nectary of two discrete
lobes nearly encircling ovary, 0.8-1 .5 x 0.7-1. 5 mm. Ovary sub-cylindrical or sub-
spherical, 0.7-1 .3 X 0.7-1 .2 mm, upper part of ovary densely pubescent to pilose, hairs
straight, to 0.1 mm long; style 3. 7-4.6 mm long, glabrous or with few straight hairs at
base; stigma c. 0.4 mm wide. Fruit sub-cylindrical or sub-spherical, 1.9-2. 8 x 2-2.5
mm; operculum 0.6-0. 9 imn, indumentum as on ovaiy. Seed narrowly ovoid to ovoid,
0.4-0. 5 X 0. 1-0.2 mm, medium to dark brown, pattern straight to spiralled, regular,
infrequently splitting and merging.

Distribution. Philippines (Mindanao).

Habitat and ecology. Lithophytic, probably on limestone in humid, shady areas.
Recorded at 83 m altitude. Flowering and fruiting March.

Rrovisional lUCN conservation assessment. Data Deficient (DD). This species is only
known from two quite old collections and the current status of the species is unknown.
The area where these collections were made is still largely forested so the population
may still be in good condition.

Additional specimen examined. PHILIPPINES: Mindanao: Agusan: Jabonga, Kitsarao, 14
Mar 1949, Mendoza, D.R. & Convocar, RR. 226 [PNH10394] (PNH).

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Notes. Epithema philippinum is most similar to E. stiigosum. See discussion under
Epithema strigosum regarding the differences between E. philippinum and E. strigosum.

Plants of Epithema philippinum that only have one leaf opposite one or more
peduncles could also be confused with E. horsfieldii but is most easily distinguished
by the straight hairs on the ovary and operculum in E. philippinum (hooked in E.
horsfieldii).

13. Epithema pusillum (C.B. Clarke) Bransgrove, stat. nov. - Epithema carnosum
var. pusillum C.B. Clarke, Monogr. Phan. 5(1): 178 (1883). - Epithema dentatum
ym. pusillum (C.B.Clarke) Hilliard & B.L.Burtt, Edinburgh J. Bot. 54: 112 (1997). -
TYPE: India, Bombay, Stocks s.n. (holotype K). (Fig. 3).

Herb 4.5-12 cm high, caulescent, indumentum of straight and hooked hairs; stem 1 mm
wide with two nodes 1.2^ cm apart and 1-8 cm to the first node, stem sub-glabrous
to sparsely pubescent, 0.2 mm long. Leaves thinly membranous, petiolate; petiole of
lowest leaf 1-2 cm long, petioles of upper leaves 0.1-1. 8 cm long, pubescent, hairs
to 0.2 mm long; blade of the lowest leaf 4.3-6 x 1 .8^.5 cm, upper leaves 2-6.5 x
1 .2^.8 cm, lowest leaf broadly ovate, upper leaves ovate, all symmetrical, apex acute
to rounded, base sub-cordate to truncate, inserted evenly on petiole, margin serrate;
upper surface weakly strigose, hairs sparse, hyaline and/or white, 0.4 nun long; lower
surface sub-glabrous to sparsely pubescent with primarily hooked hairs to 0,5 mm
long. Inflorescences 1-3 per plant; peduncles 1-2.5 cm long, usually originating from
axils of tenninal leaves, indumentum sparse, strigose, straight hairs to 0.3 mm long, or
with a sparse to dense minute hooked pubescence; bracts small and sub-cucullate to
cucullate but not enclosing entire inflorescence, 4-8 x 4-9 mm, margin sub-entire with
occasional dentation; lower surface glabrous or sub-glabrous; upper surface glabrous;
pedicels 1.9-2. 5 mm long, glabrous or with a minute, sparse pubescence. Calyx 2.1-
4.6 mm long; tube 0.8-2. 5 x 1.7-2. 5; lobes 1-2,1 x 0.7-2. 1 mm; usually glabrous or
with a minute hooked pubescence, sometimes with additional straight and/or hooked
hairs to 0.33 mm long outside; glabrous inside. Corolla white, c. 6 mm long; tube c.
4.6 mm long; lobes c. 1.4 mm long; glabrous outside, with villous band of hair inside
in upper half Stamen and staminode characters unknown. Nectary of one lobe, almost
encircling the ovary, c. 0.8 x 0.8-1. 8 mm, margin entire or sub-entire. Ovary sub-
cylindrical to sub-spherical, 0.5-1 x 0.5-1 mm, glabrous; style c. 4 mm long; stigma
c. 0.5 mm wide. Fruit sub-cylindrical, 1.5-2. 3 x 2. 1-2.3 mm; operculum c. 0.6 mm,
glabrous. Seed elliptic to ftisiform, 0.4-0. 5 x 0. 1-0.2 mm, pattern partially spiralled
to spiralled.

Distribution. India (Maharashtra).

Habitat and ecology. Lithophjdic and epiphytic (on at least Dysoxylum sp.) in humid,
probably shaded, places. Recorded from 739 m altitude. Flowering and fruiting
September.

Revision of Epithema

203

Provisional lUCN conservation assessment. Data Deficient (DD). This species is only
known from tliree collections, none of which were collected recently. The area where
these collections were made has been heavily deforested and a survey of the status of
this species is, therefore, vital.

Additional specimens examined. INDIA: Maharashtra: Ratnagiri, Amboli Ghat, Temple Point,
3 Sep 196^, Kulkami, B.G. 108673 (E); Savantwadi, 30 Sep 197^, Almeida, S.M. s.n. (E).

Notes. Epithema pusillum is easily separated from the species under which it has
been included as a variety, E. dentatum (now E. ceylanicum) and E. carnosum, by its
glabrous ovary. Epithema pusillum is only known from central, western peninsular
India. The specimen from Amboli Ghat is not as fine as the other two specimens seen
here, but the ovary and the fruit, however, are consistently glabrous.

14. Epithema rennellense Hilliard & B.L.Burtt, Edinburgh J. Bot. 54: 112 (1997). -
TYPE: Solomon Islands, Rennell Island, 18 August 1962, Dissing, H. 2738 (holotype
E; isotypes E, C).

Herb 12-27 cm high, often with an acaulescent appearance with first leaves arising
near ground level, indumentum of short pubescent hairs and longer strigose or hispid
hairs, the latter up to 0.8 mm long on petioles, upper leaf surfaces, peduncles, pedicels
and outer calyces; stem 2-4.4 mm wide with one or two nodes, intemodes 3.5-11.7
cm long. Leaves membranous to strongly membranous, petiolate, leaves solitary;
petiole of lowest leaf 7-13 cm long, upper petioles 2.5-11.5 cm long; blade of lowest
leaf 6-11.5 x 4.2-9. 3 cm, upper leaf blades 4-14 x 3.4-10 cm, all blades cordate
to oblong cordate or sub-orbicular, symmetrical or not, if asymmetrical, one side up
to 1 .4 times wider than the other, apex acute to rounded, base shallowly auriculate
to rounded, inserted evenly on petiole or slightly offset, margin dentate, bidentate
or serrate; lower surface light green or drying white-green, often finely pubescent
or hispidulous. Inflorescences 1-2 per plant; peduncles 2-21 cm long, originating
opposite single leaves or occasionally in leaf axil; bracts sub-cucullate to cucullate,
occasionally small, enclosing the entire inflorescence or portion thereof, 3-13 x 4-10
mm, margin entire with occasional crenate or dentate lobes to dentate; upper surface
sub-glabrous to sparsely pubescent, hairs to 0.2 mm long, usually on upper portion of
bract; pedicels 1 .3-5 mm long. Calyx light green, 4. 2-7. 5 mm long, tube (1 .7-)3-4.2 x
(1.5-)2-3 mm, lobes 1.5-5 x 1-1.5 mm; glabrous to sparsely strigose on lobes inside,
hairs straight or sti'aight and hooked, to 0.13 mm long. Corolla white, 10.4-15 mm
long; tube 6-11 x 2-3.3 mm; lobes 3-4.6 mm long, margin uneven/undulate; glabrous
or sub-glabrous or with a band of villous hair inside. Stamens 1. 2-3.1 mm long;
filaments 0.7-2. 1 mm long; anthers 0.5-1 mm long; staminodes 0.8-1. 5 mm long.
Nectary variable, one lobe (not entire) encircling ovary or with two discrete lobes
not encircling ovary, 0.6-1 .6 x 0.7-3. 5 mm, margin entire to undulate. Ovary 1-1 .6 x
0.8-1. 6 mm, sub-glabrous or pubescent at sparse to medium density on upper portion

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Card. Bull. Singapore 67(1) 2015

of ovary, hairs hooked or both straight and hooked, 0.04-0.3 mm long; style to 8.8
mm long, glabrous or with few hairs at base, hairs straight and hooked; stigma c. 0.5
mm wide. Fri//r obovate, cylindrical or sub-cylindrical, 2.5^ x 1-3.3 mm; operculum
0.7-1. 3 iimi, indumentum as ovary, but often on top of operculum or densest on top of
operculum. Seed nan'owly ovoid to ovoid, 0.4-0. 8 x 0. 1-0.2 mm, light to dark brown,
pattem straight to spiralled, reasonably regular, walls somewhat thickened and rigid.

Distribution. Solomon Islands (Rennell Island).

Habitat and ecology. Growing on well-drained hillsides over limestone (Rennell
Island is a limestone, coral atoll) at 20-90 m altitude.

Provisional lUCN conservation assessment. Least Concern (LC). Although this species
has not often been collected, and it is endemic to an island that is only 660 km2, the
island is still largely forested and there is little evidence that the the population is under
serious threat.

Additional specimens examined. SOLOMON ISLANDS: s.l. 1894, Officers of H.M.S.
“Penguin ” (K). Rennell Island: 21 Aug 1962, Dissing, H. 2798 (C); 6 May 1968, Sirute’e, B.
etal. BSIP9621 (K, L, SING); Hutuna, 25 Mar 1965, Wolff, T. 3004 (C); Matangi Area, 1 7 May
1969, Gafiii, I.H. & collectors BSIP14759 (K, L, SING); Nuipani, 14 Mar 1965, Wolff, T. 3025
(C); Tuhungganggo, 24 May 1969, Gafui, I.H. & collectors BSIP 14763 (K, L, SING).

Notes. This species is similar to Epithema longipetiolatum and E. strigosum in the
leaves arising at ground level. Epithema rennellense is also somewhat similar to E.
dolichopodum but differs in peduncle length/plant height ratio, leaf shape and leaf
margin, the length of the calyx and the corolla, the ovary /operculum hair and the size
of the seed. The variation in the shape of the hair on both the ovary and the operculum,
between hooked and both straight and hooked, can be seen on different fruit from the
one plant. The variation in corolla hair is unusual and is seen in few other species,
none of which are acaulescent species. Epithema rennellense has the largest seed of all
species of Epithema and, along with E. longitubum, the largest flowers.

15. Epithema sarawakense Hilliard & B.L.Burtt, Edinburgh J. Bot. 54: 113 (1997). -
TYPE: Malaysia, Sarawak, Fifth Division, Sungai Medalam, Gunung Buda, 20 June
1975, Burtt, B.L. 8336 (holotype E; isotype SAR). (Fig. 9)

Herb 25^0 cm high, sprawling or caulescent, indumentum of pubescent or strigose
and setose hairs, to 0.7 mm long on the lower leaf surface and calyx, to 1 mm long
on the upper leaf surface; stem 1.2-5 mm wide with 2 to 4 nodes, intemodes 1-8 cm
apart long. Leaves membranous to strongly membranous, petiolate, primarily solitary,
rarely with one or two sets of opposite leaves per plant; petioles 1-7 cm long; blade
2.2-17 X 1.4-13.2 cm, often cordate, but can be sub-orbicular, elliptic or sub-oblong.

Revision of Epithema

205

symmetrical or not, if asymmetrical, one side up to 1.5 times wider than the other, apex
acute to rounded, occasionally almost truncate, base usually (sub-)cordate but also to
truncate, inserted evenly on petiole or not, margin sub-entire to dentate or bidentate;
upper surface sparsely to densely strigose or villous, occasionally sub-glabrous;
lower surface may dry grey-green, glabrous to villous. Inflorescences 3-11 per plant;
peduncles 0.5-16 cm long, 1-5 arising in sequence opposite solitary leaves or rarely
in the axils of upper opposite leaves; bracts cucullate, usually completely enclosing
inflorescence, 10-32 x 6-12 mm, margin dentate or irregularly dentate; lower surface
with hairs to 0.5 mm long; upper surface often glabrous; pedicels 1.3^. 6 mm long,
often glabrous. Calyx 3. 3-6. 6 mm long; tube 1.5-3. 5 x 1-3.1 mm, lobes 1.5-3. 8 x
0.6-2. 3 mm; frequently sub-glabrous to finely pubescent outside (short, hooked hairs),
occasionally with sparse strigose hairs; inside glabrous or with few straight hairs, 0.1
mm long, in tips of lobes. Corolla white and pale blue to purple, pink, or mauve, may
have darker markings on the lobes, 8.7-11 mm long; tube 6-7 x 1.5-2. 5 mm, lobes
2. 7^. 2 X 1. 5-2.1 mm, margin slightly fimbriate. Stamens 3 mm long; filaments 1.5
mm long; anthers 1.5 mm long; staminodes 1.5-1. 8 mm long. Nectary a single lobe,
partially to completely encircling ovary, 1-1.3 x 1.2-2. 5 mm, margin almost entire to
undulate, occasionally almost dividing into smaller lobes. Ovary sub-cylindrical, 0.8-
1 X 0.7-0.9 mm, usually glabrous, occasionally ovary pubescent with hooked hairs on
upper ovary, hairs to 0.1 mm long; style c. 6 mm long, slightly wider at base, glabrous;
stigma c. 0.4 mm wide. Fruit obovate-cylindrical, 2-3 x 1 mm; operculum glabrous,
occasionally sub-glabrous, indumentum as on ovary. Seed narrowly ovoid to ovoid or
somewhat sigmoid, 0.4-0.5 x 0. 1-0.2 mm, light brown to dark brown, pattern straight
or almost so, occasionally partially spiralled to spiralled.

Distribution. Malaysia (Sabah, Sarawak), Indonesia (Kalimantan, Sumatra: Pulau
Enggano).

Habitat and ecology. Lithophytic on limestone (but substrate not always reported).
Humid, shady areas, beside streams, limestone outcrops, cave entrances. Occasionally
found on wood or on soil over rock. At 30-1250 m altitude. Flowering and fmiting
year round.

Provisional lUCN conservation assessment. Least Concern (LC). This species is
widespread with many of the collections having been made in the protected areas of
Gunung Mulu National Park and Niah National Park.

Additional specimens examined. MALAYSIA; Borneo: Sabah: Tambunan, Crocker Range,
Tambunan/Ranau Road, 5 Mar 1995, Sugau, J. JBS96 (E). Sarawak: 1st Division, Jambusan,
Gunung Batu , 6 Oct 1 977, Martin^ P.J. S. 39273 (E, K, L); Baram District, Jun 1 894, Haviland,
G.D. & Hose, C. 3525 R (K); Baram District, Batu Gading, 5 Jan 1965, van Niel, J.P. 3554
(L); Baram District, Gunung Mulu National Park, Bukit Binarat, 28 Apr 1 985, Moktar, A. et al.
S. 49467 (E, K); Baram District, Gunung Mulu National Park, Deer Cave, 16 i 1978, Hansen,
C. 17 (C, E, SAR); ibidem, 16 Jan 1978, Hansen, C. 17 (C, E); ibidem, 2 Oct 2007, Julia, S. et

206

Card. Bull. Singapore 67(1) 2015

al. S.99303 (E); ibidem, 5 May 1978, Kiew, R. RK525 (E, UPM); ibidem, 1 6 Jan 1978, Nielsen,

I. 17 (E); ibidem, 23 Jul 1987, Prirnack, R.B. S. 42 402 (K, L); Baram District, Gunung Mulu
National Park, Gua Rusa, 30 Oct 1977, Argent, G.C.G. & Kerby, R. 621 (E); Baram District,
Gunung Mulu National Park, 22 Mar 1964, Hotta, M. 15281 (E, KYO, L); ibidem, 14 Mar
1964, Hotta, M. 14402 (E, KYO); ibidem, 19 Mar 1990, Yil PC. & Abu Talib S.58618 (SAR);
Miri, Mulu National Park, en route from HQ to Deer Cave, 20 Dec 1 999, Irnaichi, R. et al. 30
(SAR); Miri, Mulu National Park, Royal Mulu Resort, 6 Oct 2001, Julaihi S. 86868 (SAR);
Miri, Mulu National Park, Simons Cave, 27 Apr 1997, Haegens, RM.A.P. & Klazenga, N. 515
(SAR); Baram District, Gunung Mulu National Park, Melinau Gorge, 23 Jun 1962, Burtt, B.L.
& Woods, P.J.B. B. 2230 (E); ibidem, 20 Feb 1978, Nielsen, 1. 416 (E); Baram District Gunung
Mulu National Park, Pala River, 14 Nov 1977, Argent, G.C.G. & Collins, M. 733 (E); Baram
District, Ulu Melinau, Aug 1958, Ashton, PS. A. 339 (K); ibidem, Aug 1958, Ashton, PS. A.
340 (K); Baram District, Ulu Sg. Tutoh, Batu Kalulong, 13 Apr 1997, Julaihi et al. S. 76938
(SAR); Batu Niah, Nov 1 932, Synge, PM. S.565 (K); ibidem, Nov 1 932, Synge, PM. S.634 (K);
Batu Niah, Gunung Subis, 28 Nov 1966, Anderson, J.A.R. et al. S. 26090 (A, E, K, L, SING);
ibidem, Jan 1961, Mohidin S.21608 (K); ibidem, 19 Aug 2002, Sabli, J. et al. S. 89048 (SAR);
Batu Niah, Niah Cave Park, 4 Jun 1962, Burtt, B.L. & Woods, P.J.B. B.2003 (E); ibidem, 3
Jun 1962, Chew, W.-L. CWL.293 (K, L, SAR, SING); ibidem, 23 Dec 1999, Irnaichi, R. et al.
38 (SAR); ibidem, 20 Aug 2006, Prieditis, N s.n. (E); ibidem, 20 Aug 2006, Prieditis, N. s.n.
(E); ibidem, 18 Apr 1978, Stone, B.C. 13730 (K, KLU, L); Belaga District, Bukit Merirai,
Gua Pak Danum, 12 Jul 2005, Leong, P. et al. PL320 (SING); Belaga District, Ulu Merirai, 9
Jul 2005, Leong, P. et al. PL215 (SING); Belaga District Ulu Merirai, Gua Tiang, 6 Jul 2005,
Julia, S. et al. S.93376 (SING); Bintulu, Ulu Sg. Kakus, Bukit Sarang, 14 Mar 1965, Anderson,

J. A.R. S. 2095 7 (A, K); Binmlu, Ulu Sg. Kakus, Bukit Sarang, Batu Anyi, 5 Oct 2004, Julia,
S. et al. S. 9475 7 (SAR, SING); Buseau, 1890, Haviland, G.D. s.n. (K); Kapit, Melmau, 16
Jul 1961, Anderson, J.A.R. & Keng, H. KlOO (SAR); Kapit, Ulu Melmau, Gunung Api, 1971,
Anderson, J.A.R. S.31770 (E, K, L, SING); ibidem, 10 Sep 1970, Chai, P. S.30372 (A, BO, K,
L, SING); ibidem, 1970, Lehmann, P.F. PPL. 596/S, 30372 (E); ibidem, 7 Sep 1970, Lehmann,
PE S.30091 (E, K, SAR); ibidem, 29 Mar 1990, Yii, PC. & Abu Talib S.58841 (SAR); ibidem,
29 Mar 1990, Yii, PC. & Abu Talib S.58841 (SAR); Lobang Rusa, 9 Jun 1975, Burtt, B.L. B.
8227 (E, SAR); Lobang Rusa, Sungei Melinau Paku, 9 Feb 1966, Chew, W.-L. CWLlOl 7 (A, K,
L, SING); Marudi District The Pinnacles, Gunung Api, 17 Jun 1995, Beaman, J.H. 11727 (K);
Miri, Gunong Mulu National park. Hidden Valley, 5 Apr Argent, G.C.G. et al. 905b (E);
Niah, 7 Oct 1954, Ahmad I (SING); Niah National Park, 12 Dec 1981, Rogstad, S.H. 728 (A).
INDONESIA: Sumatra: Bengkulu: Pulau Enggano, Boea-boea, 11 Jun 1936, Lutjeharms,
W.J. 4598 (L); Pulau Enggano, Malakoni, 26 Jun 1936, Liltjehaims, W.J. 5112 (BO, GH, K, L,
P). Kalimantan: Peningin, Jo/zm 1522 (BO).

Notes. Epithema sarawakense is usually quite distinctive with its near complete lack
of opposite leaves and one to five infloresences arising from the base of the solitary
petiole. Epithema sarawakense is often densely hairy, giving the dried leaves a grey-
green colour. There are two forms, however, one of which is glabrous or sub-glabrous.
The glabrous form is found almost exclusively in Gunung Mulu National Park in the
environs of Deer Cave and the base of Gunung Mulu. While there are a small number
of collections of the pubescent fonn from near Deer Cave, it has primarily been
collected from Gunung Api, Gunung Buda, Gunung Benarat, along Sungai Melinau
and in the Melinau Gorge.

Revision of Epithema

207

There are a small number of specimens that show some similarity to Epithema
saxatile. These specimens have some opposite leaves but, in most cases, there are
always solitary leaves with peduncles arising opposite the solitary leaf on the plant.
These opposite leaves are unequal in size while Epithema saxatile has even-sized
opposite leaves. In addition, the range of the corolla length of Epithema saraM^akense
is 8-11 mm while that of E. saxatile is 6. 5-9. 5 mm and the upper o vary/ operculum
of E. sarawakense is glabrous or sub-glabrous (hairs hooked) while the operculum
of E. saxatile is always at least sparsely pubescent (hairs hooked). The specimens of
Epithema sarawakense that are similar to E. saxatile are not segregated geographically
from more typical E. sarawakense.

A small number of specimens from Pulau Enggano in Sumatra are included in
Epithema sarawakense. While the distribution is difficult to explain, there are currently
no characters to separate the specimens from the remainder of the species as found in
Sarawak. They have shorter stems than is usual for Epithema sarawakense, thereby
resembling E. strigosum, but they differ from E. strigosum in that the operculum is
glabrous to sparsely pubescent with hooked hairs rather than dense and with straight
hairs as in E. strigosum.

16 . Epithema saxatile Blume, Bijdr. 738 (1826); Ridley, FI. Malay Penins. 2: 539
(1923); B.L.Burtt, Thai For. Bull. (Bot.) 29: 94 (2001). - TYPE: Indonesia, Java,
Blume, C.L. 1029 (lectotype L [L0003232], designated here; probable isolectotype U
[without number]). (Fig. 11, 12)

Epithema carnosum auct. non Benth.: Barnett, FI. Siam. 3(3): 205 (1962), p.p.

Herb 6-40 cm high, caulescent; stem 2-5 mm wide with 2-5 nodes, intemodes
1.2-10.5 cm long, usually glabrous or sub-glabrous. Leaves membranous to strongly
membranous, petiolate or with sessile upper leaves, rarely with one or two solitary
upper leaves; petiole of lowest leaf 2.5-12 cm long, upper petioles 0-2.5 cm long;
blade oflowest leaf 7-20 x 6.5-12.5 cm, upper leaves 4.5-1 3.5 x 3-8.5 cm, lowest leaf
sub-spherical to broadly ovate, upper leaves variable, often narrowly ovate to almost
oblong, but also cordate, broadly ovate or elliptic, symmetrical or not, if asymmetrical,
one side up to 1.4 times wider than the other, apex acute to broadly acute, rarely
rounded, base cordate to truncate, inserted evenly on petiole to distinctly offset (up to
2 cm), margin sub-entire to dentate, bidentate or serrate (often all or degrees of these
on one plant); upper surface green or dark-green, indumentum occasionally to 1.1 mm
long; lower surface light green, whitish or occasionally purplish. Inflorescences 1-10
per plant; peduncles 2-17 cm long, indumentum occasionally hispid, to 0.7 mm long;
bracts cucullate, usually enclosing the entire inflorescence, 13-35 x 8-12 mm, margin
usually dentate, occasionally sub-entire with one or two teeth or lobes; lower surface
usually sparsely to densely strigose, hairs straight or primarily straight, 0. 3-0.5 mm
long; upper surface glabrous or sub-glabrous, hairs towards margins of bract; pedicels
1.1^.2 mm long. Calyx 3.3-5 mm long; tube 1.6-3. 3 x 1. 5-2.1 mm; lobes 1.5-2. 7 x

208

Card. Bull Singapore 67(1) 2015

Fig. 11. Epithema saxatile Blume. Flowers and cucullate bract. (Photo: David Middleton)

Fig. 12. Distribution of Epithema saxatile Blume (•).

Revision of Epithema

209

0.6-0. 9 nmi, often glabrous at maturity, hairs to 0.5 mm long, glabrous inside. Corolla
white, pink, blue, purple, or purple and white, frequently with highly variable purple
markings on the upper lip, 6. 5-9.5 mm long; tube 5-7 x 1.5-2. 5 mm; lobes 1-3 x
1.7-2. 3 mm; band of villous hair up to 1 mm wide inside. Stamen filaments 1. 3-2.1
mm long; anthers c. 0.6 mm long; staminodes 1. 3-2.1 mm long. Nectary apparently
absent or with one or two lobes partially to entirely encircling ovary, margin undulate,
0.7-1. 3 X 0.6-2 mm. Ovary sub-cylindrical or sub-obovate, 0.6-1. 5 x 0.5-1 mm, very
sparsely to densely pubescent, hairs hooked, on the upper or very upper portion of
ovary, hairs 0.04-0.63 nnu long; style 2.9-5 mm, glabrous or sparsely pubescent on
the lower style; stigma 0.3 mm wide. Fruit cylindrical or obovate, 1.9-2. 6 x 1.7-2. 1
mm; operculum 0.4-0. 7 mm long, pubescence as ovary. Seed narrowly ovoid to ovoid,
0.3-0. 5 X 0. 1-0.2 mm, pattern stright to spiralled, reasonably regular.

Distribution. Myanmar, Thailand, Malaysia (Peninsular Malaysia, Sarawak, Sabah),
Indonesia (Sumatra, Java, Kalimantan, Sulawesi).

Habitat and ecology. Shaded, humid areas, usually on or amongst limestone rocks or
in limestone soil, near, by or in streams or rivers. Also found near or on the walls of
cave mouth entrances. At 0-910 m altitude. Flowering and fruiting hugely variable.

Provisional lUCN conservation assessment. Least Concern (LC). This species is
common and widespread.

Additional specimens examined. MYANMAR: Tenasserim, 1877, Gallatly, G. 1035 (SING);
Tenasserim, Tavoy, Paungdaw, Aug 1 96\, Keenan, J. et al. 798 (E).

THAILAND: Chanthaburi: Nam Tok Phliu National Park, Nam Tok Phliu Waterfall, 30
Aug 1969, Maxwell, J.F s.n. (AAU); Priu Waterfall, 12 Oct 1965, Chermsirivathana, C. 413
(BKF). Chiang Mai: Doi Sutep, 4 Aug 1912, Kerr, A.F.G. 2655 (K); Me Ta Chang, 17 Oct
1922, Ken; A.F.G. 6381 (ABD, E, K). Chon Buri: Bahn Beung District, Ang Chang Nam, 1
Sep 1975, Maxwell, J.F. 75-958 (AAU, BKF, L); Pong NamRawn, 3 Sep 1956, Smitinand, T.
3489 (BKF, E); Sriricha, Chundaten Falls, 1 7 Aug 1 974, Maxwell, J.F. 74-814 (AAU, BK, L),
Chumphon: Bang Son, 11 Sep 1927, Put 1051 (ABD, K). Kanchanaburi: Between Huay
Ban Kao and Kritee, 4 Jul 1973, Geesink, R. & Phengklai, C. 6082 (BKF, E, L); ibidem, 12 Jul
1973, Geesink, R. & Phengklai, C. 6231 (AALT, BKF, C, E, K, L, P). Kanchanaburi: Hindato,
24 Jul 1946, Kosterrnans, A.J.G.H. 1346 (BO, A, L, P); Dongyai, 14 Aug 1971, Phengklai, C.
2933 (BKF); Kliaibuing, CP. BS& 5A 2997 (BKF, E, L); Kin Sayok, 13 Jul \946,Kostermans,
A.J.G.H. 1103 (A, BO, L, P, SING, US); Lai Wo, Toong Yai Naresuan Wildlife Reserve, Ban
Saneh Pawng, 12 Aug 1993, Maxwell, J.F. 93-886 (CMU); Rintin, 31 Jul 1946, Kosterrnans,
A.J.G.H. 1401 (A, BO, L, P, SING, US); Si Sawat, Hua Lum Kao Ngoo, 5 Jul 1 973, Sutheeson,
S. 2489 (BKF); Tawng Pa Poom, Huay Ban Kliao, 12 Jul 1973, Maxwell, J.F 73-285 (AAU).
Khainphaeng Phet: Khlong Lan National Park, 21 Aug 1995, Parnell, J. et al. 95-335 (K).
Krabi: 29 Mar 1930, Kerr, A.F.G. 19385 (K); Ao Luk, 21 Jun 2006, Williams, K. et al. 2033
(A); Nai Chong, 21 Oct 1979, Sutheeson, S. 5004 (BKF). Mae Hong Son: Muang, Nahng
Rawng Falls, 16 Sep 1972, Maxwell, J.F. 72-362 (L). Nakhon Nayok: Nang Rong Falls, 13
Aug 1968, Larsen, K. et al. 3363 (AAU, BKF, C, E, K, L, P, SING). Nakhon Si Thammarat:
Thung Song District, Khao Tham Long, 31 Aug 1982, Shimizu, T. et al. T. 28985 (BKF); Nop

210

Card. Bull. Singapore 67(1) 2015

Phitum, Khao Luang National Park, Krung Ching Falls, 24 Sep 2010, Middleton, D.J. et al.
5524 (E); ibidem, 12 Feb 2005, Williams, K. et al. 1400 (E). Phangnga: Khao Lak, 21 Sep
1963, Smitinand, T & Sleumer, H.O. 1207 (BKF, L). Phangnga: Muang, Suan Somdet, 8 Dec
1999, Wongprasert, T, 9912-30 (BKF); Muang, Tham Pha Phueng, 15 Sep 2010, Middleton,
D.J. etal. 5425 (E); Panga, Sep 1894, Curtis, C. s.n. (SING); ibidem, Aug 1893, Curtis, C. s.n.
(SING); Pulau Tebun, 29 Dec 1918, Nur, M. 3585 (K, SING); Talcuapah, 14 Jul 1972, Larsen,
K. et ai 30967 (AAU, E); Tham Tong Lang, Shimizu, T, et al. Z29151 (BKF). Phatthalung:
Khao Pu-Khao Ya National Park , 24 Sep 1986, Maxwell, J.F. 86-723 (A, BKF, L, PSU).
Phitsanulok: N. Phitsanulok, Thung Salaeng Luang National Park, 22 Jul 1966, Larsen, K.
et al. 722 (AAU, BKF, L); ibidem, 25 Jul 1973, Murata, G. et al. T.17092 (AAU, BKF, KYO,
L); ibidem, Phusomsaeng, S. et al. 83 (BKF); ibidem, 30 Sep 1967, Tagawa, M. et al. T. 11 259
(KYO); Thung Salaeng Luang National Park, Kaeng Sopa Waterfall, 22 Oct 1984, Murata, G.
et al. 38544 (BKF); ibidem, 22 Oct 1984, Murata, G. et al. T. 38467 (BKF); Thung Salaeng
Luang National Park, Kang So Pa Waterfall, 17 Sep 1990, Chantharanothai, P. et al. 90/303
(K); Thung Salaeng Luang National Park, Poi Waterfall, 22 Oct 1984, Murata, G. etal. T.38587
(BKF). Saraburi: Khao Muak Lek, 2 Oct 1963, Bin Rajab, M.K. 714 (KLU). Songkhla:
Boripath Waterfall, 1990, Larsen, K. et al. 41243 (AAU); ibidem, 16 Aug 1984, Maxwell, J.F.
84-55 (A, PSU); Boriphat Falls National Park, 9 Nov 1990, Larsen, K. et al. 41243 (P). Surat
Thani: Ban Kawp Kiep, 5 Aug 1927, Kerr, A.F.G. 13183 (ABD, E, K). Surat Thani: Klilong
Phanom National Park, Middleton, D.J. et al. 4345 (BKF, E); Phanom, Khao Sok National
Park, 6 Sep 2008, Middleton, D.J. et al. 4330 (E); Phanom, Khao Sok National Park, Chong
Lorn, 12 Dec 1979, Shimizu, T. et al. T.27103 (KYO); ibidem, 12 Dec 1979, Shimizu, T. et al.
T.27111 (BKF, KYO); ibidem, 12 Dec 1979, Shimizu, T. etal. T.27100 (KYO); Phanom, Khao
Sok National Park, Sii Ru Cave, 27 Feb 2006, Middleton, D.J. et al. 4055 (BKF, E); Sawng Pi
Nawng, 20 Mar 1927, Kerr, A.F.G. 12385 (K). Satun: Khuan Don, Thale Ban National Park,
Along cliff face from Ton Din Cave, 9 Sep 2010, Middleton, D.J. et al. 5350 (E); Thale Ban
National Park, 27 Aug 1995, Larsen, K. et al. 46055 (AAU); Thung Wa, Than Plew Waterfall,
1 0 Sep 2010, Middleton, D.J. et al 5368 (E). Trang: Ampoe Kao Kao, Kao Chom Lem, 2 Aug
1929, Rabil 318 (K). Trang: Ban Nam Phrai, 17 Dec 1979, Shimizu, T. et al T.27551 (BKF,
KYO, L); ibidem, 17 Dec 1979, Shimizu, T. etal T.27571 (KYO); Chawng, 15 Mar 1959,
Smitinand, T. & Abbe, E.C. 6149 (BKF, K); Huay Yot, Wat Tham Iso, Middleton, D.J. et al.
4428 (E, SING); Lamphura, 15 Nov 1990, Larsen, K. et al. 41413 (AAU, BKF); ibidem, 16
Nov 1990, Larsen, K. et al. 41450 (AAU); Trang, Nam Tai, 1 1 Oct 1970, Charoenphol, C. et
al. 3662 (AAU, E); ibidem, 1 1 Oct 1970, Charoenphol, C. et al. 3661 (AAU, BKF); Trat, Koh
Chang, 7 Sep 1992, Niyomdham, C. 3276 (BKF). Yala: Betong, 27 Aug 1923, Kerr, A.F.G.
7687 (K).

LAOS: Chanipassak: Thorel, C. s.n. (P); ibidem, Thorel, C. s.n. (P).

VIETNAM: Luang, Aug 1 868, Pierre, L. 4542 (P). Dong Nai: Bien Hoa, Cochinchine, Pierre,
J.B.L. 4541 (P); Gia Lai: Dak Doa, Poilane, E. 18108 (P).

MALAYSIA: Sabah: 24 Feb 1985, Lamb, A.L. 8/85 (E); Interior Zone, Batu Punggol, Apr 1987,
Venneulen, J.J. 1188 (L); Kinabatangan District, Kori Timber Camp, 7 Nov 1948, Cuadra, A.
A. 2168 (A, K, KEP, SAN, SING); Kinabatangan District, Northern Tabin Wildlife Reserve
(Near Tabin River), 16 Oct 2000, Poulsen, A.D. et al. 1659 (E); Lahad Datu, Bukit Baturong, 7
Jul 2000, Kiew, R. RK5040 (SING); Lahad Datu, Tabin Wildlife Reserve, 24 Oct 2000, Kiew, R.
RK5120 (SING); ibidem, 25 Oct 2000, Pius, G. etal. SANl 43649 (E, K); Madai Baturong Forest
Reserve, Baturong Hill, 12 Jim 1996, Lim, S.P. et al. LSP738 (SING); Sandakan, Gomantong,
13 Feb 1960, Meijer, W. SAN20756 (A, K, L); Sandakan, Gomantong, Gomantong Caves, 31
Oct 1968, Kokawa, S. & Hotta, M. 551 (KYO); ibidem, 31 Oct 1968, Kokawa, S. & Hotta,

Revision of Epithema

211

M. 557 (KYO); ibidem, 31 Oct 1968, Kokawa, S. & Hotta, M. 565 (KYO); ibidem, 25 Sep
1968-26 Sep 1968, Ogata, K. 10975 (KYO, L); Sandakan, Gomantong, Gomantong Hill, 20
Sep 1970, Banggilon, K. SAN66579 (K, L); ibidem, 28 Apr 1996, Lirn, S.P et ai 580 (SING);
ibidem, 26 Apr 84, Sands, MJ.S. & Young, R.G.N. 3936 (E, K); Sandakan, Hutan Simpan
Gomanton, Between Gomantong Cave and Bukit Dulong Lambu, 10 Sep 1976, Taniura, M.
& Hotta, M. 602 (KYO); Sandakan, Tabin River, 16 Oct 2000, Venneulen, J.J. 2037 (SING);
Sukau, Panggi, 18 Sep 1996, Kiew. R. & Urn, S.R RK4211 (K, SING); Tawao, Oct 1922-Mar
1923, Elmer, A.D.E. 20569 (G, GH, K, SING). Sarawak: 1st Division, 28 May 1975, Biirtt,
B.L. B.8207 (E); 1st Division, Ban, 24 Apr 1955, Brooke, W.M.A. 9872 (L); 1st Division, Ban
District, Burtt, B.L. & Woods, P.J.B. B. 1898 (E); 1st Division, Ban District, Bukit Jebong, 7 Aug
1970, Lehmann, PF PFL364 (E); 1st Division, Bau District, Bukit Kapor, 22 May 1962, Burtt,
B.L. & Woods, P.J.B. B.1887 (E); 1st Division, Bau District, Fairy Cave, 9 Feb ]999, Jemree,
S. et al. S. 82056 (SAR); 1st Division, Bau District, Seburan Mine, 24 May 1962, Burtt, B.L. &
Woods, P.J.B. B.1925 (E); 1st Division, Bau Limestone Hills, Bukit Boring, 14 Nov 1985, Yii
et ai S. 503 5 3 (K, L); 1st Division, Bau, Guniing Stulang, 16 Oct 200\, Meekiong, K. SBC778
(SING); 1st Division, Bulit Majing, Tebakang, 16 Dec 1988, Kessler 226 (L); 1st Division,
Gunung Berloban, 19 Apr 1984, Yii & Othman S.46225 (K, L); 1st Division, Jambusan, Poak
Road, 16 Dec 1989, Frodin, D.G. & Bin Ismawi, O. 2077 (K); 1st Division, Padawan, Bukit
Megetang, 4 Mar 1969, Wright, E. & Chai, P S. 27 4 5 7 (E, K); 1st Division, Padawan, Gunung
Regu, 6 May 1975, Burtt, B.L. B.8089 (E); 1st Division, Taiton, Yii et al. S. 5 1 207 (L, SAR);
Bidi, Jul 1893, Ridley, H.N. s.n. (SING); Bidi Cave, Oct 1929, Clemens, J. & Clemens, M.S.
7434 (NY); ibidem, 22 Oct 1929, Clemens, J. & Clemens, M.S. 7627 (K); Kuching, Idzurni, H.
(Sc Togashi, M. s.n. (TI); ibidem, 29 Oct 1975-31 Oct \91 5, Idzurni, H. (Sc Togashi, M. s.n. (TI);
Kuching to Padawan, 30 Sep 1981, Croat, T.B. 53161 (SAR); Kuching, Daerah Kecil Siburan,
Kampong Mambong, 12 Dec 1999, Imaichi, R. et al. 2 (SAR); Kuching, Pemissen, Gunung
Braang, 8 Dec 2000, Jemree, S. etal. S. 84036 (SAR); Kuching, Penrissen, Mambong, Gunung
Bar, 19 Feb 1993, Lai & Rantai et al. S. 66009 (K, SAR); Kuching, Tiang Bekap, Mt. Mentawa,
1 1 Mar 1 967, CheM’, W.-L. CWL. 1288 (A, AAU, K, L, SING); Kuching, Tiang Bekap, Padawan
Road, 3 Apr 1960, Anderson, J.A.R. 12349 (K, L, SING); Samarahan , Sedan District, Lobang
Mawang, Bukit Selabor, 26 Sep 1968, Ilias bin Pale S. 28047 (K, L, SING); ibidem, 26 Sep
1968, Ilias bin Paie S. 28051 (K); Tambusan, Sep 1904, Ridley, H.N. s.n. (SING); Mongkos,
Gunong Selebur, 20 Apr 1999, Jamree et al S. 82239 (SAR); Sedan Distinct, Gunung Niyat,
Ulu Sg. Majat, 27 Feb 2002, Jemree, S. & Enjah, A. S.85584 (SAR). Peninsular Malaysia:
Kedah: Baling, 5 Aug 1941, Naven, J.C. 38038 (SING); Kota Setar, G. Keriang, 24 Jul 2009,
Rosdi, M, FRI66329 (SING); Pulau Langkawi, Gn. Raya Forest Reserve, 5 Oct 2010, Chan,
Y.M. FRI70632 (SING); Pulau Langkawi, Selat Panchor, 23 Nov 1934, Henderson, M.R.
29079 (K, SING); Langkawi, 20 Nov 1941, Corner, E.J.H. s.n. (SING); ibidem, 20 Nov 1941,
Corner, E.J.H. s.n. (SING); ibidem, Nov 1889, Curtis, C. 2107^.^ (SING); ibidem, Curtis, C.
2107a (SING); Langkawi, Gunung Raya, 22 Aug \9'^^, Anthony samy, S. SA959 (SING); Pulau
Langkawi, Pulau Dayang Buting, 23 Aug 1925, Holttum, R.E. 15125 (BO); Langkawi, Pulau
Langgun, 4 Nov 1971, Chin, S.C. 1759 (KLU). Kelantan: Bukit Mangga, 20 Feb 2003, Kie^^

R. RK5276 (SING); Gua Maka, 19 May 1990, Kiew, R. (S Anthonysamy, S. RK3020 (SING);
Kuala Aring, 1 Sep 1992-11 Sep 1992, Hamid H18 (SING). Pahang: Bukit Cheras, 10 Oct
1901, Henderson, M.R. 25205 (NY, SING); ibidem, 26 Nov 1984, Kiew, R. RK1566 (UPM);
Gunung Jebak Puyuh, Kiew, R. RK2145 (UPM); Gunung Senyum, 22 Aug 1986, Anthonysamy,

S. SA515 (UPM); Kampong Sri Jaya, Bukit Batu, 28 Oct 1986, Kiew, R. RK2369 (SING);
Kuala Lipis, 25 Aug 2008, Mohd Hairul, M.A. FRI60060 (SING); Kuantan, Bt Cheras, 25
Aug 1986, Anthonysamy, S. SA558 (UPM); Kuantan, Bukit Punching, 26 Nov 1984, Kiew’, R.

212

Card. Bull. Singapore 67(1) 2015

RK1570 (UPM); Maran, Jengka Forest Reserve, Hutan Lipur Jebak Puyuh area, 14 Oct 2008,
Mohd Hairul, M.A. FRI60066 (SING); Taman Negara, Batu Kepayang, 3 Oct 1984, Kiew, R.
RK1423 (UPM); Taman Negara, Batu Luas, 1 Oct 1984, Kiew, R. RK1349 (UPM); Temerluh,
Gn. Jebak Puyuh, Gn. Senyum, 9 Feb 2011, Kamarul Hisham, M, FRI67230 (SING). Perak:
Jan 1885, Kings Collector 7046 (SING); ibidem, Nov 1880, Kiinstler, H. 983 (SING); ibidem,
1890, Ridley, H.N. s.n. (SING); Batu Kurau, Dec 1884, Scortechini 1580 (SING); Chemar,
Perak Tong Temple, 23 Oct 1958, Sinclair, J. 9846 (E, L, SING, US); Grik, Flutan Simpan
Kemerong limestone area, 18 Nov 2011, Mohd Hairul, M.A. FRI 54071 (SING); Ipoh, 1 Dec
1966, Ng, FS.R FRI1795 (L, SING); Ipoh, Gunung Rapat, 2 Mar 1959, Allen, B.M. 4272
(SING); ibidem, 14 Jan 1983, Davis 69304 (E); Kampong Jahang, Gunung Kanda, 5 May
1962, Biirtt, B.L. & Woods, P.J.B. B.1820 (E); Larut, Apr 1884, King’s Collector 5872 (SING);
Larut & Matang, Lata Puteh, 22 Nov 2008, Yao, T.L. FRI65393 (SING); Tambun, Rotan Segar,
29 Nov 1960, Allen, B.M. 4654 (SING). Perils: Besih Hangat, 17 Nov 1929, Henderson,
M.R. 22875 (SING); Bukit Bintang Forest Resei've, Bukit Bintang, 9 Aug 1986, Weber, A.
860809-1/1 (E); Mata Ayer F.R., Biildt Rongkit, Kiew, R. RK3712 (KEP); Wang Kelian, 28 Jun
1993, Kiew, R. RK3653 (SING). Selangor: Kuala Lumpur, Jul 1890, Curtis, C. s.n. (SING);
Batu Caves, 24 Aug 1908, Ridley, H.N. 13380 (K, SING); ibidem, 4 Sep 1966, Burkill, H.M.
HMB4223 (SING); ibidem, 10 Jul 1906, Ernst, A. 1118 (L); ibidem, 23 Jan 1966, Hardial, S. &
Sidek474 (C, K, L, SING); ibidem, 9 Jun 1999, Kmv, R. RK4711 (SING); ibidem, 15 Sep 1968,
Kokawa, S. 6381 (KYO); ibidem, Dec 1898, Ridley, H.N. 8217 (SING); ibidem, Dec 1891,
Ridley, H.N. s.n. (SING); ibidem, 30 Oct 1967, Shimizu, T. & Fukuoka, N. M. 14 143 (AAU,

K, KYO, L, SING); Genting Highlands, Ulu Gombak, 26 Oct 1937, Nur, M. 34253 (SING).
Terengganu: Batu Biwa, 22 Oct 1986, Kiew, R. RK2287 (SING).

INDONESIA: Java: s.l., 1857-1861, de Vriese, W.H. s.n. (L); Unknown 2648 (L); Gunung
Buning (Gunung Bunder), 19 Dec 1893, Schiffner, V.F. 2567 (A, BO, K, L); Gombak, Sempor,
17 Apr 1936, Brinkman, R. 672 (BO). Jawa Timur: s.l., Coert, J.H. 1093 (L); Kangean Island,
Ardjasa, 1920, Backer, C.A.B. 26969 (BO); Kangean Island, Batoe Poetih, 28 Mar 1919,
Backer, C.A.B, 27805 (BO); Kangean Island, Tambajangan, 22 Mar 1919, Backer, C.A.B. 27410
(BO); Soekapoera, 3 Jun 1927, Backer, C.A.B. s.n. (BO); Tretes, Gunung Ai'djoeno, May 1919,
Bremekamp, C.E.B. s.n. (BO); Zuidergebergte, 27 Mar 1927, Backer, C.A.B. & Posthumus,
O. s.n. (BO); Madioen, Ngebel, 16 Mar 1898, Koorders, S.H. 29801 (K, L); Pogal, Mousset
450 (BO, L). Jawa Tengah: Lebak Barang, 12 Jan 1918, Backer, C.A.B. 23287 (BO). Jawa
Barat: Pasir Masigit, Feb 1933, Jacobson, E. 213 (BO); Papandajan, Mar 1930, van der Pijl,

L. 205 (BO); Bandung, Zollinger, H. 2002 (G, GH, P, S); Bogor, 1909, Backer, C.A.B. s.n. (L);
Bogor, Gunung Tjibodas , Feb 1912, Backer, C.A.B. 2480 (L); Cibodas, 1910, Backer, C.A.B.
32476 (BO); ibidem, 1917, Backer, C.A.B. 22091 (BO); ibidem, 1917, Backer, C.A.B. 22091
(BO); ibidem, 18 Mar 1928, van Steenis, C.G.G.J. 616 (BO); Tjiampea, Gunung Tjibodas,
15 Feb 1924, Bakhuizen van den Brink, R.C. 3568 (BO, L); ibidem, 30 Jan 1921, Bakhuizen
van den Brink, R.C. 5153 (BO, L); Klappa Noenggal, 1912, Backer, C.A.B. 5848 (BO, L);
Padelarang, Kampong Sempang, Coert, J.H. 625 (L); Res. Preanger, Tjidadap, Tjibeber, 23
Feb 1917, Bakhuizen 2608 (L); Tjampea, Burck, W. & de Monchy, B.J. s.n. (BO); ibidem,
9 Feb 1913, Koorders, S.H. 40438 (BO); ibidem, de Monchy, B.J. s.n. (L); ibidem, 4 May
1895, Hallier, H.G. 91 (L); ibidem, 25 Sep 1904, Hochreutiner, B.P.G. 1916 (G); UJung Kulon
National Park, Tjilintang, 25 Apr 1963, Wirawan, N. 45 (A, BO, K, L, SING); Palimena, Mont.
Prope, Junghuhn, F.W. 3 (K, L). Sumatra: s.l., Yates, H.S. 1939 (NY). Aceh: Gajolanden, 25
Feb 1937, van Steenis, C.G.G.J. 9267 (BO, K, L); Gaju and Alas Lands, Pendeng to Bivouac
Aer Putih Waterfall, 17 Feb 1937, van Steenis, C.G.G.J. 8879 (BO, L); Gunung Leuser Nature
Reserve, Ketambe Research Station, Gunung Giring, 17 Mar 2008, Wilkie, P. etal. PW764 (E);

Revision of Epithema

213

Gunung Leaser Nature Reserve, Ketambe, Mt Ketambe, 24 May 1972, de Wilde, W.J.J.O. & de
Wilde- Duyjjes, B.E.E. 12353 (BO, K, L); Ketambe, 17 Mar 2008, Sumadijaya, A. AX360 (BO);
Kloet Nature Reserve, South Kloet, Pucuk Lembang, 10 Jul 1985, de Wilde, W.J.J.O, d de
Wilde- Duyjjes, B.E.E. 19894 (BO, L); Pulau Weh, Sabaug, 31 Dec 1933, van Steenis, C.G.G.J.
5734 (BO). Lampung: Gunung Rati, 24 Nov 1921, Unknown 208 (BO, L); Muaradua, 13 Apr
1929, de Voogd, C.N.A. 357 (BO, L). Riau: Anambas Islands, Siantan Island, Terempak, 6 Apr
1928, Henderson, M.R. 20277 (K, SING). Sumatera Barat: Jorong Gasang, Lake Maninjau,
18 Jun 2{)\\ , Puglisi, C. & Hughes, M. CP61 (BO); Padang, Indarung, Kanipong Putih, 16 Feb
1981, Hotta, M. & Okada, H. 247 (KYO); Pajakumbuh, Halaban, 2 Feb 1958, Maradji 482
(L, SING). Sumatera Selatan: Aer Telanai, 11 Nov 1929, Van Steenis, C.G.G.J. 3925 (BO).
Sumatera Utara: Pulau Tello, Nov 1924, Boden-Kloss, C. s.n. (SING); Batu Islands, Pualu
Telo, 23 Jan 1 897, Unknown 654 (BO); Upper Langkat, 6 Nov 1 938, Lorzing, J.A. 1 7359 (BO);
Wampu, 1918-1919, Lesger 359 (BO). Borneo: Kalimantan Timur: West Koetai, Kombeng,
22 Nov 1925, Endert, F.H. 5143 (BO, L); West Koetai, Lahren, 29 Jun 1925, Endert, F.H. 1762
(BO, L); Tanah Grogot, Batu Kajang, Desa Kasungai, 27 Nov 1979, Ma ’wef, A. AM.250 (BO,

K) ; Berau, Sungai Kelai, Gunung Njapa, 18 Oct 1963, Kostemians, A.J.G.H. 21338 (BO, G, K,

L) ; Berau, Tanjung Redeb, Gmiung Buntung, 28 Nov 198 1-1 Sep 1981, Kato, M. et al. B. 11 733
(BO, KYO); Kenangan, 12 Aug 1979, Dransfield, J. 4402 (BO). Kalimantan Selatan: 1908,
Winkler, H. 2670 (BO, G, K, L); Gunung Serempaka, 26 Nov 1971, Dransfield, J. & Saerudin,
D. 2314 (BO, L); Djaro Dam, 11 Nov 1971, Kuswata 718 (BO); Muara Uja, 21 Nov 1971,
Dransfield, J. & Saerudin, D. 2268 (BO, K, L). Sulawesi Selatan: Bantimurong National
Park, 20 Feb 1938, Buwalda, P 3761 (BO), Sulawesi Tengah: G. Batoe, 1913, Rachmat 439
(BO); Luwuk, 9 Oct 1989, Coode, M.J.E. 5855 (K); Soroako-Wasuponda Road, 7 Jul 1979,
van Balgooy, M.M.J. 3922 (L). Sulawesi Tenggarah; Pohara, Muara Sampara, 14 Apr 1929,
Kjellberg, G.K. 1299 (BO, S); Pulau Butung, Jismal Camp, 12 Nov 1989, Coode, M.J.E.
6206B (K); Pulau Butung, 17 Feb 1929, Kjellberg, G.K. 195 (S). Sulawesi Utara: Bogani
Nani Waitabone National Park, 24 May 2002, Uji, T. 4595 (BO).

Notes. Epithema saxatile is one of the most widely distributed species of Epithema.
At first it appears that there are a few different forms of E. saxatile, one with petiolate
upper leaves, one with sessile upper leaves and one with occasional, solitary upper
leaves, but all of these can be found within a single population.

Some collections of this species have strongly variegated leaves but this
character does not appear to have any taxonomic significance.

17. Epithema steenisii Hilliard & B.L.Burtt, Edinburgh J. Bot. 54: 113(1 997). - TYPE:
Indonesia, Sumatra, Aceh, Gaju & Alas Lands, Gadjah to Pendeng, 16 February 1937,
van Steenis, C.G.G.J. 8825 (holotype L; isotypes BO (image seen), K, SING). (Fig.
10 )

Herb 3-1 5 cm high, caulescent, indumentum of strigose, setose or hispid hairs; stem
1-2 mm wide with 2-4 nodes, intemodes 1-3 cm long. Leaves strongly membranous,
petiolate, occasionally with more than one solitary lower leaf; petioles 0.2^. 5 cm
long; blade of lowest leaf 3.2-9 x 1 .7-4 cm, upper leaves 1 .5-9 x 0.7-4. 7 cm, more or
less lanceolate, elliptic, or naiTowly ovate, upper portion sometimes curved and upper

214

Card. Bull. Singapore 67(1) 2015

halves asymmetric, if the blade is asymmetrical, one side up to 1.5 times wider than
the other, apex acute to more rarely broadly acute, rarely rounded, base sub-truncate,
more rarely cuneate to rounded, inserted evenly on petiole or not, margin variable,
weakly serrate to bidentate; upper surface densely strigose to villous, hairs 0.75-1
mm long; lower surface densely pubescent or strigose and setose, hairs to 1 . 1 mm
long. Inflorescences 2^ per plant; peduncles 0.5-10.5 cm long, arising from leaf
axils; bracts cucullate, often completely enclosing inflorescence, 9-23 x 5-12 mm,
margin dentate, teeth small, lower surface strigose, hairs to 0.6 mm long, upper surface
glabrous or sub-glabrous; pedicels 1.5-3. 6 mm long, often with only strigose hairs
to 0.5 mm long, but sometimes also with additional minute hairs. Calyx 3.5-5 mm
long; tube 2-3 x 1 .2-2.3 mm, 1-2 times as long as wide, lobes 1. 5-2.7 x 0.6-1. 5 mm;
outside sparsely to densely strigose, hairs straight, occasionally hooked, 0.5-0.8 mm;
glabrous inside. Corolla white or pale lilac to lilac, 6.7-9 mm long; tube 4-6.5 x 1-1.5
mm; lobes 2.5 x 1 mm. Stamen filaments c. 1.2 mm long; anther characters unknown;
staminodes c. 1.2 mm long. Nectary of two discrete lobes, c. 0.8 mm high. Ovary
sub-cylindrical, c. 0.9 x 0.8 mm, sub-glabrous to densely pubescent, hairs hooked,
0.05 mm long and placed on upper portion of ovary; style 3. 7-4. 6 mm long, glabrous
or sub-glabrous, hairs at base of style. Fruit obovate to cylindrical, 1.4— 2.3 x 1.7-
2.3 mm; operculum 0.5-0. 8 mm, sub-glabrous to sparsely pubescent, as ovai*y. Seed
sub-cylindrical or narrowly to broadly ovoid, the ends often only slightly constricted,
0.3-0. 6 X 0. 1-0.3 mm, dark brown, pattern straight to spiralled and reasonably regular.

Distribution. Indonesia (Aceh).

Habitat and ecology. Shaded areas, on and among wet (mossy) rocks and in wet sand
by streams and rivers at 200-600 m altitude. Flowering and fruiting February, March,
May, June, August.

Provisional lUCN consei^ation assessment. Vulnerable VU D2. This species is only
known from the Indonesian province of Aceh in northern Sumatra and many of the
populations are from Gunung Leuser National Park. The species is only known to
occur below 600 m altitude and Gunung Leuser has been subjected to continued
deforestation despite being a “protected” area (Barber et al., 2002).

Additional specimens examined. INDONESIA: Aceh: Between Lau Simerah and Lau
Penanggajan, 20 Mar 1954, Alston, A.H.G. 14535 (A); Gaju & Alas Lands, Pendeng-Bivouac
Aer Putih Waterfall, 17 Feb 1937, van Steenis, C.G.G.J. 8879A (BO, L); Gunung Leuser Nature
Reseiwe, Camp Simpang, 18 Aug 1972, de Wilde, W.J.J.O. & de Wilde- Duyfles, B.E.E. 14353
(BO, K, L, US); Gunung Leuser Nature Reserve, Lau Alas Valley, Ketambe, 21 May 1972, de
Wilde, W.J.J.O. & de Wilde- Duyfjes, B.E.E. 12238 (BO, L); Ketambe, Alas River, 9 Jun 1979,
de Wilde, W.J.J.O. & de Wilde-Duyjjes, B.E.E. 18042 (BO, K, L).

Notes. This species is very similar to Epithema tenerum from Sulawesi but can be
distinguished by the difference in the indumentum on the outside of the ealyx. In

Revision of Epithema

215

addition the cucullate bract usually encloses the inflorescence in Epithema steenisii
whereas this is variable in E. tenerum. See further discussion in the notes section of
Epithema tenerum.

18. Epithema strigosum (C.B. Clarke) Hilliard & B.L.Burtt, Edinburgh J. Bot. 54: 113
(1997). - Epithema brunonis var. strigosum C.B.Clarke in A.DC. & C.DC., Monogr.
Phan. 5(1): 179. (1883). - TYPE: Indonesia, Sumatra, Gunung Singgalan, Korthals,
RW. 692 (lectotype L, designated by Hilliard & Burtt (1997) [see also note below]).
(Fig. 10)

Herb 6-20 cm high, acaulescent or with first leaves arising from a very short stem;
stem 2-4 nun wide with 0-1 nodes, intemodes 0-1.5 cm long. Leaves membranous to
strongly membranous, petiolate, one lower solitai^ leaf and one upper, or several leaves
in short-stemmed tuft; petioles 0.5-21 cm long; blade 1-19 x 0.5-15 cm, cordate to
elongate ovate, symmetiical or not, apex acute, base cordate, sub-auriculate or rounded,
inserted evenly on petiole or not, margin crenate or dentate, teeth wide at base and
mostly with rounded tips; both surfaces with hairs to 0.8 mm long. Inflorescences 1^
per plant; peduncles 6.5-23.5 cm long, originating opposite solitary leaves or at base
of petioles; bracts cucullate, completely enclosing the inflorescence, 14-18 x 8-15
mm, margin entire to dentate, lower surface setose and strigose, hairs medium density,
to 0.4 mm long, upper surface glabrous to strigose, hairs to medium density, primarily
straight or straight and hooked, to 0.4 mm long; pedicels 1 .7-5(-7.5) mm long, finely
densely pubescent, hairs hooked, also with or without sparse straight hairs 0. 1-0.4 mm
long. Calyx 3. 1-5.2 mm long; tube 1. 7-4.2 x 1.5-2. 7 mm, lobes L5-2.3 x 0.8-1. 7
mm; sparsely setose and strigose outside, hairs primarily hooked or both straight and
hooked, white, to 0.4 mm long, covering calyx; inside with hairs throughout to only on
lobes, hairs straight or both straight and hooked, to 0.13 mm long. Corolla white, 4.5-
8.3 mm long; tube 3.7-5 x 1.5 mm; lobes 0.8-3. 3 mm long; margin slightly fimbriate;
a partial or complete band of villous hairs in upper half inside. Stamens 2. 1-2.4 mm
long; filaments 1 .5-1 .8 mm long; anthers c. 0.6 mm long; staminodes 1 .2-1 .7 mm long.
Nectary of one or two lobes, 0. 5-0.3 mm long. Ovary 0.8-1 x 0.7-0. 8 mm, densely
pilose, hairs straight, white, 0. 1-0.3 mm long, placed on upper portion of ovary; style
3. 5^. 6 mm long, glabrous or sparsely pubescent at base of style, hairs straight; stigma
c. 0.5 mm wide. Fruit cylindrical to spherical, 1 .5-1 .9 x 1 ,1-2.3 mm; operculum c. 0.6
mm long, pubescence as on ovary. Seed ovoid, sometimes somewhat curved, 0.4-0.5
X c. 0. 1 mm, light to medium brown, pattern almost straight to spiralled, regular.

Distribution. Indonesia (Sumatra).

Habitat and ecology. Lithophytic. Found in shaded, humid areas beside rivers and
streams and along paths at 100-1000 m altitude. Flowering and fruiting January-
March, August.

216

Card. Bull. Singapore 67(1) 2015

Provisional lUCN conservation assessment. Endangered (EN Blab(iii)). This species
has a restricted distribution in West Sumatra in areas subject to continuing degradation
of natural habitats.

Additional specimens examined. INDONESIA: Sumatera Barat: KabupatenPadangPariaman,
Sipisang, 9 Jan 1995, Okada, H. et al. 1343 (BO); Gunung Singgalan, Singalang, Korthals,
P.W s.n. (L); Mt Sago, 9 Mar 1957, Meijer, W 5637 (L); Padang, Indarung, K. Putih, 16 Feb
1981, Hotta, M, Okada, H. & Kohyama, T. 319 (KYO); Padang, Indai’ung, Ladang Padi, K.
Putih, 8 Aug 1984, Hotta, M. & Okada, H. 1087 (KYO); ibidem, 14 Aug 1981, Hotta, M. &
Okada, H. 516 (KYO); Padang, Ladang Padi, 28 Aug 1984, Hotta, M., Okada, H. & Kohyama,
T. 689 (KYO, L); Padang, Ulu Gadut, 1 0 Jan 1 983, Hotta, M., Okada, H. & Ito, M. 402 (KYO);
Padang, Ulu Gadut, Sungai Gadut Gadang, 2 Aug 1984, Hotta, M., Okada, H. & Kohyama, T
324 (KYO); ibidem, 2 Aug 1984, Hotta, M., Okada, H. & Kohyama, T 339 (KYO).

Notes. Epithema strigosum is most similar to E. philippinum but differs from E.
philippinum in the larger calyx, the lack of a minute pubescence on the outside of
the calyx and the larger hairs only being sparse, the pubescence inside the calyx is
less dense, the leaf margin is crenate rather than dentate or serrate, the corolla can be
longer (4.5-8. 3 imn long rather than 5-6 mm long) and is reportedly white rather than
blue. Epithema strigosum is only found in Sumatra and E. philippinum only in the
Philippines. All specimens of Epithema strigosum are from an area of limestone east
of Padang.

Korthals s.n. (L0796748) is possibly a duplicate of Korthals 692, the type
collection, but has no field number, only a herbarium number of 113 which is also on
the lectotype. The plants are very similar and could be the same collection.

19. Epithema tenerum (C.B. Clarke) Hilliard & B.L.Burtt, Edinburgh J. Bot. 54: 113
(1997). - Epithema brunonis var. tenerum C.B. Clarke in A.DC. & C.DC., Monogr.
Phan. 5(1): 180 (1883). - TYPE: Indonesia, Sulawesi, South Sulawesi, Maros,
Zollinger, H. 1172 (lectotype G, designated here; isolectotypes L, P). (Fig. 6, 13)

Herb 5-14(-20) cm high, caulescent; stem 0.8-2 mm wide with 2-3 nodes, intemodes
0.5-5(-l 0) cm long. Leaves thinly membranous to membranous, petiolate, upper leaves
opposite or sub-opposite; petiole of lowest leaf 1-5 cm long, upper petioles 0.2-5 cm
long. Blade of the lowest leaf 2.3-12 x 1.4-8 cm, upper leaves 1.9-10 x 0.9-5 cm,
leaves usually broadly lanceolate to narrowly ovate or elliptic, but occasionally broadly
ovate or obovate to broadly obovate, symmetrical or not, if asymmetrical one side up
to 1 .4 times wider than the other, apex acute to broadly acute, base sub-cordate, sub-
tmncate or obtuse, inserted evenly on petiole or not, margin entire to weakly serrate;
upper surface primarily strigose, hairs straight, medium density, to 0.7 mm long; lower
surface may dry grey-green, sub-glabrous to pubescent, hairs medium density, to 0.4
mm long, veins raised with distinct, sparsely to densely pubescent, primarily strigose
hairs, to 0.6 mm long. Inflorescences 1-6 per plant; peduncles 0.3-7(-18) cm long,
originating from the terminal and lower axils; bracts small, sub-cucullate or cucullate.

Revision of Epithema

111

Fig. 13. Epithema tenerum (C.B. Clarke) Hilliard & B.L.Burtt. A. Habit. B. Underside of leaf.
C. Inflorescenee. D. Calyx. E. Flower opened out. F. Stigma lateral view. G. Fruit showing
seeds, placenta and operculum. H. Seeds. Scale bars; A = 4 cm; B = 5 mm; C, E = 3 mm; D, G
= 2 mm; F = 1 mm; H = 0.5 imn. Drawn by Claire Banks from Zollinger 1172 (A), Zollinger
1064 (B), P.C. Thomas and W H. Aroli 09-74 (C-G), Jaag 1600 (H).

218

Card. Bull. Singapore 67(1) 2015

6-16 X 3-7 mm, imdiilate to lobed with occasional dentation or dentate; lower surface
strigose, hairs straight, to medium density and 0.5 mm long; upper surface glabrous
or sub-glabrous; pedicels 0.8-3, 3 mm long, finely pubescent hairs to 0.13 mm long,
also with or without straight, strigose hairs to 0.5 mm long. Calyx occasionally tinged
purple, 1.7-3 .4 mm long; tube 0.9-2. 5 x 0.9-2. 5 mm; lobes 0.6-1. 5 x 0.6-1. 9 mm;
sub-glabrous, strigose or setose outside, also with minute, hooked hairs, hairs medium
density, straight hairs to 0.5 mm long, hooked hairs to 0.3 mm long; glabrous inside.
Corolla blue, pale-blue or pale purple, 5.2-6 mm long, tube 4-4.8 x 0.8-1. 3 mm;
lobes 0.9-1. 8 x 1-2.1 mm, margins entire to slightly crenate, dense band of villous
hair inside. Stamen filaments c. 1.5 mm long; anther characters unknown; staminodes
c.1.25 mm long. Nectary usually of two discrete lobes (occasionally one), margin
entire, 1 x 0.75-1.5 mm. Ovary sub-cylindrical, 0. 8-0.9 x 0.6-0.75 mm, with hooked
hair on upper half of ovary, hairs to densely pubescent, c. 0.1 mm long; style c. 3.3
mm long, glabrous; stigma up to c. 0.5 mm wide. Fruit sub-spherical, 1. 6-2.1 x 1.9-
2.2 mm, operculum 0.6-0. 8 mm long, indumentum as ovary. Seed narrowly ovoid to
ovoid, 0.4-0. 5 x 0. 1-0.2 mm, light to medium brown, pattern sub-erect to spiralled,
reasonably even.

Distribution. Indonesia (Sulawesi).

Habitat and ecology. Lithophytic, often found directly on limestone or on rocks of
unknown type at 20-750 m altitude. Usually beside streams and waterfalls but also on
road sides and on rock walls.

Provisional lUCN conservation assessment. Vulnerable (VU D2). This species is only
known from a small number of localities and these mostly karst limestone, a habitat
which is frequently exploited for cement and tourism.

Additional specimens examined. INDONESIA: Sulawesi Utara: Bolaang Mongondow,
Dumoga Bone National Park, Edwards Camp, 26 Mar 1985, de Vogel, E.F & Vermeulen, J.J.
6759 (L); Maros, Zollinger, H. 1064 (A, P). Sulawesi Selatan: Bantimurung, 16 Jun 1938,
Jaag, 0.1600 (L); ibidem, 20 May 1929, Rant, A. 40 (BO); ibidem, 20 Feb 1938, Buwalda,
P 3671 (L); ibidem, 20 Feb 1938, Biiwalda, P. 3677 (BO, L); ibidem, 13 Apr 1975, Meijer,
W.9133 (BO, L, US); Bantimurung, Bantimurung Waterfall, 15 Apr 2009, Thomas, D. &Ardi,
W.H. 09-74 (E); Pankadjene, 8 May 1931, Teruya, Z. 1848 (SING). Sulawesi Tenggara:
Kesali-Porema, 23 Oct \929, Kjellherg, G.K. 2618 (BO, S).

Notes. This species is most similar to Epithema steenisii from Aceh, Sumatra. It differs
mainly in the type of calyx hair, the density of the leaf hair and the length of the
corolla. On the calyx there are small, hooked hairs in addition to the longer, straight
hairs which are present in both Epithema tenerum and E. steenisii. In addition, the
longer hairs are generally shorter in Epithema tenerum than in E. steenisii (to 0.5 mm
vs. to 0.8 mm long). The hair on the upper surface of the leaves is usually much less
dense in Epithema tenerum than in E. steenisii and the corolla tube is 4—4.8 nmi and
4-6.5 mm respectively.

Revision of Epithema

219

Epithema tenerum usually follows the general Epithema pattern of having one
larger, lower leaf and one or more pairs of opposite, upper leaves. Some upper leaf
pairs are actually sub-opposite and this is most clearly seen in the larger specimens
(eg. Teruya 1848 (SING)).

The species has been collected from few localities in Sulawesi, principally from
Bantimurung in the south-west of the island. The specimens from northern Sulawesi,
De Vogel and Vermeulen 6759 (L) and, in particular, Kjellberg 2618 (S), are generally
larger than those from Bantimumng. The multi-specimen sheet from Pankadjene,
Teruya 1848 (SING), in southern Sulawesi, however, has plants spamiing the entire
size range.

20 . Epithema tenue C.B.Clarke in A.DC. & C.DC., Monogr. Phan. 5(1): 181 (1883). -
TYPE: Equatorial Guinea, Bioko, 1863, Mann, G. 2345 (lectotype K, designated here;
isolectotype P). (Fig. 14, 15)

Epithema thomense Henriq., Bol. Soc. Brot. 10: 145 (1892). - TYPE: Sao Tome and
Principe, Sao Tome, May 1888, Quintas, F 1272 (lectotype COI, designated here;
isolectotype K).

Epithema graniticolum A.Chev., Bull. Soc. Bot. France 58 (Mem. 8d.): 189 (1912).
- TYPE: Guinea, Montagne de Boola, Pays de Guerzes, 1048 m, 16 March 1909,
Chevalier, AJ.B. 20924 (lectotype P, designated here).

Herb 5-40 cm higli, acaulescent or caulescent, occasionally the plant is of only one leaf;
stem 1-5 mm wide with 1-3 nodes, intemodes 1-16.5 cm long. Leaves membranous
or strongly membranous, lower leaves petiolate, upper leaves petiolate and/or sessile
or occasionally with a solitary, upper leaf opposite peduncles; petiole of lowest leaf
2-10 cm long, upper petioles 0.2-8 cm long; blade of lowest leaf 6.5-24 x 4.5-19
cm, upper leaves 2-10 x 1. 5-8,4 cm, the lowest leaf very large and almost oblong to
broadly ovate, ovate or cordate, upper leaves ovate to broadly elliptic or spherical,
symmetrical or not, apex often rounded but also acute, base cordate or sub-cordate,
inserted evenly on petiole or base of upper leaves with one side truncate or obtuse
and the other cordate, margin variable, often sub-entire or minutely serrate or dentate,
but also serrate, dentate or crenate (usually the upper leaves); upper surface weakly
strigose or strigose, hairs to medium density, to 1.5 mm long; lower surface strigose
to medium density, 0. 5-0.8 mm long, and pubescent with minute, hooked hairs to
0.1 mm long. Inflorescences 1-6 per plant; peduncles 0.5-14 cm long, originating
from the base of the stem, opposite the solitary lower leaf or from the tenninal axil;
bracts frequently very large, cucullate and enclosing the inflorescence but may be
small and enclosing a small proportion of the inflorescence, 4-40 x 5-14 mm, margin
dentate or entire; lower surface weakly strigose or strigose, hairs 0.2-0. 8 mm long;
upper surface glabrous or sub-glabrous, hairs straight and concentrated towards
margins; pedicels 0.8^ mm long, weakly strigose, hairs to medium density and 0.8

Fig. 14. Epithema tenue C.B. Clarke. A. Habit. B. Inflorescence. C. Calyx. D. Flower opened
out. E. Stigma lateral view. F. Fruit showing seeds, placenta and operculum. G. Seeds. Scale
bars: A = 1 0 cm; B = 5 mm; C, F = 3 mm; D = 6 mm; E = 1 mm; G = 0.5 mm. Drawn by Claire
Banks from Letouzy 7722 (A, B), Letouzy 13973 (D, E) and Sita 2886 (C, F, G).

Revision of Epithema

221

Fig. 15. Distribution of Epithema tenue C.B. Clarke (•).

mm long, also hooked hairs to medium density and 0.1 mm long. Calyx 2-8 x 1.3-2
mm; tube 1.3-4 mm long; lobes 0.8-3. 8 mm long; with long hairs usually villous, or
occasionally to glabrous or sub-glabrous in maturity, hairs to high density, especially
in young flowers, hyaline, 0.6-1 .5 mm long; glabrous inside. Corolla blue-purple, tube
white, 9.6-10,5 mm long; tube 6. 6-7. 5 x 2 mm; lobes c. 3 mm long; a band of villous
hairs inside, hairs at half to two-thirds the way up the tube. Stamens 2-2.5 mm long;
filaments 1.5-2 mm long; anthers c. 0.5 mm long; staminodes 1.5-2 mm long. Nectary
usually apparently absent, but also of one or two lobes, sometimes reduced, 0.6-1 x
0.8-1. 3 mm, margin undulate or entire. Ovary ovoid, spherical or cylindrical, 0.8-1. 5
X 0.6-1. 3 mm, weakly hirsute to villous with hairs straight, 0.5-1. 5 mm long, with or
without additional hooked hairs to 0. 1 mm long (often seen most easily on the fruit
operculum); style 6 mm long, glabrous or one or two hairs at base, hair to 0.08 mm
long; stigma c. 0.5 mm wide. Frw/7 sub-cylindrical to sub-spherical, 1.5-3 x 1 .5-3 mm;
operculum 0.5-1 mm long, indumentum as on ovary. Seed nan’owly ovoid to ovoid,
0.4-0, 5 X 0. 1-0.2 mm, medium brown, pattern sub-erect or erect, reasonably even.

Distribution. Guinea, Sierra Leone, Liberia, Ivory Coast, Nigeria, Cameroon,
Equatorial Guinea, Gabon, Sao Tome and Principe, Democratic Republic of Congo,
Uganda, South Sudan.

Habitat and ecology^ Primarily lithophytic, often on granite at 180-1100 m altitude.
Found in humid areas, typically in shade, usually on or near rocky areas, in or beside
streams, waterfalls or cave entrances. Also found beside paths and highways. Flowering
and fruiting year round.

222

Card. Bull. Singapore 67(1) 2015

Provisional lUCN conservation assessment. Least Concern (LC). This is a widespread
species not known to be under any immediate tlireats.

Additional specimens examined. GUINEA: Kindia, Jacqiies-Felix, H. s.n. (P).

SIERRA LEONE: Between Kumbonla and Seredu, 1 5 Nov 1 965, Morton, J.K. SL2530 (E, K);
Loma Mountains, Oct 1944, Jaeger, P. 227 (K).

LIBERIA: Kolahun, Vahon, 7 Nov 1947, Baldwin, J.T. 10215 (K, US).

IVORY COAST: Chevalier, A.J.B. 21418 (K, P); Chevalier, A.J.B. 21506 (P); Banco, 11 Apr
1979, Knedl, M. 721 (G); Gueoule, 3 Feb \ 9S4, Hepper, F.N. & Maley, J. 8073 (K); Haut Nuon,
Chevalier, A.J.B. 21141 (P); Haut Sassandra, Droupleu to Zoanle, 5 May 1909, Chevalier,
A.J.B, 21457 (P); Man, Mt Tankoui, 4 Jul 1966, Ake Assi, L. 9043 (G); ibidem, Aug 1978,
Knedl, M. 9/365 (G); Tiapleu, 25 Sep 1955, Unknown 3296 (P).

NIGERIA: Gongola, Mambilla, River Nwum Forest Reseiwe, 30 Jun 1976, Chapman, J.D.
4505 (K); Sarduna, Gembu, Kamatan Forest Reserve, 14 Dec 68, Daramola, B.O. FHI62313
(K).

CAMEROON: Bamenda Province, Bamenda District, Metschum Falls, 26 Aug 1952, Savory,
H.J. UCI299 (K); Bamenda Province, Wum, Nbika’s By-Pass, 28 Jun 1951, Ujor, E. FHI29260
(K); Banyo, Tako-Atta Manga, 5 Jul 1967, Letouzey, R. 8732 (P); Collines a 5km au SW
d’Ebianeme-Yong pres Nyabessan, 10 Apr 1968, Letouzey R. 9238 (P); Ga Village, Pagan
Hill, 29 Aug 1 966, Letouzey, R. 7722 (COI, K, P); LolodorL 1 5 Jun 1918, Annet, E. 33 7 (K, P);
Muyuka, Munyenge, 29 May 1 976, Letouzey, R. 15020 (P); Muyuka, Munyenge, Mt Cameroun,
29 May 1976, Letouzey, R. 15026 (K, P); Nkambe, Vallen Forest, 12 Nov 1974, Letouzey, R.
13198 (P); Northern Cameroon, Munkep, Essu-Munkep Trail, 7 Jul 1975, Letouzey, R. 13973
(P); South West Province, Etinde, Upper Boando, Upper Boando Village, 1 Dec 1993, Cable,
S. 283 (K); South West Province, Fako, Buea, Likombe, 2 Sep 1992, Nkeng, P. 90 (K); South
West Province, Kupe-Muanenguba Division, Kupe Village, 27 May 1996, Etuge, M. 1973 (K);
ibidem, 10 Jul 1996, Kenfack, D. 259 (K); ibidem, 15 Jul 1996, Zapfack, L, 929 (K); South
West Province, Kupe-Muanenguba Division, Ngomboku, 10 Dec 1999, Mackinder, B.A. 303
(K); South West Province, Kupe-Muanenguba Division, Nyale, 17 Nov 1988, Cheek, M.R.
et al. 9647 (K); South West Province, Kupe-Muanenguba Division, Nyasoso, 24 Jun 1996,
Cable, S, 3256 (K, WAG, YA); ibidem, 19 Dec 1993, Cable, S. & Ajebe, E.F 644 (K); ibidem,
2 Jun 1996, Etuge, M. 2057 (K); ibidem, 19 Oct 1995, Sidwell, K. et al. 305 (K); South West
Province, Kupe-Muanenguba Division, Nyasoso, Mount Kupe, 7 Feb 1996, Cable, S. 3532 (K,
YA); ibidem, 7 Oct 1998, Harris, D. 5815 (E); ibidem, 23 Oct 1995, Sebsebe Demissew 4984
(K); ibidem, 7 Jul 1992, Wheatley, J.I. 4117 (K); ibidem, 2 Jun 1996, Zapfack, L. 597 (K);
South West Province, Meme, Nyasoso, 23 Jun 1996, Cable, S. 3208 (K); South West Province,
Meme, Nyasoso, Mt Kupe, 5 Aug 1993, Balding, S. & Sivell, D. 37 (K); ibidem, 19 Sep 1992,
Cable, S. 19 (K).

EQUATORIAL GUINEA: Bioko, 16 Jan 1947, Guinea, E. 1408 (K); Bioko, Basile, 30 Aug
1990, Carvalho 4464 (K); ibidem, 29 Jul 1986, Carvalho 2130 (K).

GABON: 13 Mar 1925, Le Te.stu, M.G. 5449 (P); Lastoursville, 1929-1931, Le Testu, M.G.
7018 (K); Libreville, 2 Feb 1 899, Klaine, R.P 151 7 (K); ibidem, Klaine, R.P. 2666 (P); Ndende,
22 Nov 1983, de Wilde-Dujfjes, B.E.E. et al. 745 (K).

SAO TOME AND PRINCIPE: St Thome, Henriques, J.A. 5 (K); ibidem, Henriques, J.A. 12
(K).

DEMOCRATIC REPUBLIC OF CONGO: Meya, Camp O.R.S.T.O.M., 12 Feb 1969, Sita, P.
2886 (P).

Revision of Epithema

223

UGANDA: 1906, Bagshawe, A.W.G. 1282 (US); Bundibugyo, Sempaya Hot Springs, 14 Sep
1997, Lye, K.A. & Katende, A.B. LYE22885 (K); Bwamba, Balanga, 22 Nov 1935, Thomas,
A.S. ThJ529 (K); Bwamba, Bulanga, 30 Sep 1932, Thomas, A,S. Th.729 (K); Lake Albert
Edward, 1906-1907, Bagshawe, A. W.G. 1376 (US).

SOUTH SUDAN: Imatong Mountains, Talanga, 26 Nov 1980, Friis, I. & Vollesen, K.B. 462
(C, K).

Notes. There are two forms of Epithema tenue, an acaulescent and a caulescent form.
The acaulescent fonn is found in the Central African Republic, Equatorial Guinea
(Bioko), Cameroon and Gabon while the caulescent fonn is found in the Ivory Coast,
Nigeria, Sudan, Uganda, Liberia, Guinea, Sierra Leone, Cameroon and Gabon. In
only Cameroon and Gabon are there collections of both fonns. The acaulescent form
has, primarily, big bracts and inflorescences and calyces which are pubescent with
long hairs. The size of the inflorescence is more variable in the caulescent form, but
can be still large and pubescent. The duplicates of Harris 5815 (E) from Cameroon
show the acaulescent and caulescent forms can be found within one population. In this
collection the acaulescent specimens are small and membranous and have small bracts
and inflorescences but still have the characteristic long, straight hair on the ovary/
operculum.

Some of the caulescent specimens could be mistaken for Epithema carnosum.
They differ IroniE. carnosum by the size of the bract and calyx and the dense indumentum
on the outside of the calyx. Some, however, have small bracts, inflorescences and
calyces, and the indumentum is sparse. These can still be separated using the length of
the hair on the operculum. While this hair is straight in both species, in Epithema tenue
it is from 0.5-1 .5 mm long and in E. carnosum it is from 0.08-0.25 mm long. There
are a small number of specimens that are not readily differentiated from Epithema
carnosum as the indumentum on the ovary is short. While it is sparse, the indumentum
of the calyx of these specimens is that of E. tenue, being hyaline, villous and long
instead the hyaline to white, short and more strigose hair of E. carnosum. As these
two species occur on different continents confusion between the two should not be a
problem.

Excluded Species

Epithema triandrum (Blanco) Fem.-Vill., Nov App. 150. (1880). - Ophiorrhiza
triandra Blanco, FI. Filip., ed. 2 [F.M. Blanco] 65 (1845). = Ophiorrhiza oblongifolia
DC. (see Merrill, 1918).

ACKNOWLEDGEMENTS. We thank the Sibbald Trust for providing the funding for this
revision. We thanlc the directors and curatorial staff of the herbaria that loaned material or
hosted visits. Martin Pullan is gratefully acknowledged for his help with the Padme taxonomic
database. Claire Banks is thanked for the beautiful illustrations and Serena Lee is thanked

224

Card. Bull. Singapore 67(1) 2015

for her work on the maps. Junji Matsumura (FU) is thanked for providing images of the type
of Epithema calcicola and Abdulrokhman Kartonegoro (BO) for images of isotypes of E.
horsfieldii van epiphyllum, E. longitubum and E. steenisii. The staff of the Forest Herbarium
Bangkok and Pramote Triboun are gratefully aeknowledged for enabling and assisting field
work in Thailand for the second author. Hamiah Atkins and Ruth Kiew are thanked for their
thorough reviews of the manuscript.

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Appendix 1. Index of exsiccatae.

Epithema benthamii C.B, Clarke (1); Epithema carnosum Benth. (2); Epithema ceylanicum
Gardner (3); Epithema dolichopodum Hilliard & B.L.Burtt (4); Epithema horsfieldii (R.Br.)
A.DC. (5); Epithema involucratum (Roxb.) B.L.Burtt (6); Epithema longipetiolatum
(Merr.) Hilliard & B.L.Burtt (7); Epithema longitubum Hilliard & B.L.Burtt (8); Epithema
madulidii Hilliard & B.L.Burtt (9); Epithema membranaceum (King) Kiew (10); Epithema
parvibracteatum Hilliard & B.L.Buitt (11); Epithema philippinum (Hilliard & B.L.Burtt)
Bransgrove (12); Epithema pusillum (C.B. Clarke) Bransgrove (13); Epithema rennellense
Hilliard & B.L.Burtt (14); Epithema sarawakense Hilliard & B.L.Burtt (15); Epithema saxatile
Blume (16); Epithema steenisii Hilliard & B.L.Burtt (17); Epithema strigosum (C.B. Clarke)
Hilliard & B.L.Burtt ( 1 8); Epithema tenerum (C.B. Clarke) Hilliard & B.L.Burtt (19); Epithema
tenue C.B.Clarke in A.DC. & C.DC. (20).

Aet & Idjan 32 (7), 942 (7); Ahmad 1 (15); Ake Assi, L, 9043 (20); Allen, B.M. 4272 (16),
4654 (16); Almeida, S.M. s.n. (13); Alston, A.H.G. 14535 (17); Anderson, J.A.R. et al.
S.26090 (15); Anderson, J.A.R. 12349 (16), S.20957 (15), S.31770 (15); Anderson,
J.A.R. & Keng, H. KlOO {15)-, Annet, E. 337 (20); Anthonysamy, S. SB7 (4); SA515 (16),
SA558 (16); SA959 (16); Aigent, G.C.G. et al. 905b (15), 29038 (1); Argent, G.C.G.
C8754a (7); Argent, G.C.G. & Collins, M. 733 (15); Argent, G.C.G. & Kerby, R. 621

(15) ; Ashton, RS. A.339 {\5),A.340 {\5)\Atha, D. 4785 (2).

Backer, C.A.B. s.n. (16), 2480 (16), 2789 (5), 5848 (16), 6607 (5), 6687 (5), 11469 (5), 22091

(16) , 23287 (16), 26969 (16), 27410 (16), 27805 (16), 32476 (16), 36621 (5); Backer,

C. A.B. & Posthumus, O. s.n. (16); Bagshawe, A.W.G. 1282 (20), 1376 (20); Bakhuizen
van den Brink, R.C. 2608 (16), 3568 (16), 5153 (16); Balalaishnan, N.P. 10815 (2);
Balding, S. & Sivell, D. 37 (20); Baldwin, JET. 10215 (20); Banggilon, K. SAN66579
(16); Barnes, E. 928 (3), 1056 (3), 1300 (3); Bartlett, H.H. 7740 (1 0), 14487 (3), 14744
(3), 15380 (3); Beaman, J.H. 10068 (4), 11727 (15); Beumee, J.G.B. 1328 (5); Bin
Rajab, M.K. 714 (16); Bloembergen, S. 3458 (8), 3496 (8); Blume, C.L. s.n. (16), 1029
(16); Boden-Kloss, C. s.n. (16); Boerlage, J.G. 528 (7); Brahmam, M. 2776 (3); Brandis,

D. 676 (3); Bremekamp, C.E.B. s.n. (16); Brinkman, R. 672 {\6y, Brooke, W.M.A. 9872
(16); Brown, R. s.n. (6); Biinnemeijer, H.A.B. 10713 (5), 10890 (5); Burck, W. & de
Monchy, B.J. s.n. (16); Burkill, H.M. HMB4223 (16); Burtt, B.L. B.8089 (16), B.8207
(16), B.8227 (15), B.8336 (15); Burtt, B.L. & Woods, P.J.B, B.1820 (16), B.1887 (16),
B.1898 (16), B.1925{\6\ B.2003{\5\ B.2230 {\5)\ Buwalda, P. 3671 (19), 3677(19),
3761 (16), 5110 (7), 5202 (7), 5748 (7), 6006 (7), 7225 (5), 7542 (5).

Cable, S. 19 (20), 283 (20), 3208 (20), 3256 (20), 3532 (20); Cable, S. & AJebe, E.F 644
(20); Carvalho 2130 (20), 4464 (20); Chai, P. S.30372 (15); Chan, Y.M. FRI70632 (16);
Chantharanothai, P. et cd. 90/303 (16); Chapman, J.D. 4505 (20); Charoenphol, C. et

226

Card. Bull. Singapore 67(1) 2015

al. 3661 (16), 3662 (16), 3976 (10); Cheek, M.R. et al. 9647 (20); Chennsirivathana,

C. 1 (3), 413 (16); ChevaHer, A.J.B. 20924 (20), 21141 (20), 21418 (20), 21457 (20),
21506 (20); Chew, W-L. CWL.293 (15), CWLJ017 (15), CWL.1288 (16); Chin, S.C.
392 (11), 1404 (10), 1514 (10), 1759 (16); Clung, R.C 6421 (2); Clarke, C.B. 8997 (2),
12300 (2), 15665 (2), 19076 (2), 24805 (2), 36192 (2); Clason, E. W. C99 (5); Clemens,
J. & Clemens^ M.S. 7434 (16), 7627(16); Clemens, M.S. 724 (1); Coert, J.H. 625 (16),
1093 (16); Coode, MJ.E. 5313 (1), 5855 (16), 6206B (16); Corner, EJ,H s.m (16);
Coster SM. (5); CP, BS & BN 2997 (16); Croat, T.B. 53161 (16); Ctiadra, A. A.2168
(16); Cuming, H. 823 (3), 1265 (1); Curtis, C. s.n. (16), 955 (10), 2107 {16), 2107a (16).

Daramola, B.O. FHI62313 (20); Davis 69304 (16); de Monchy, B.J. s.n. (16); de Vogel, E.F.
& Vermeulen, J.J. 6759 (19); de Voogd, C.NA. 357 (16); de Vriese, W.H, s.n. (16); de
mide, J.J.FE. 1155 (1); de Wilde, W.J.J.O. & de Wilde- Duyfjes, B.E.E. 12238 (17),
12353 (16), 14353 (17), 18042 (17), 19894 (16), 21918 (5), 21940 (5); de Wilde-Dujfjes,
B.E.E. et al. 745 (20); Deschamps s.n. (3); Dhal, N.K. & Rout, N.C. 8122 (3); Dissing,
FI. 2738 (14), 2798 (14); Dransfield, J. 3869 (1), 4402 (16); Dransfield, J. & Saerudin,

D. 2268 (16), 2314 (16); Duthie, J.F. 2990 (2).

Eberhardt 3363 (3); Edaho, G. s.n. (1), 418 (1), 435 (1), 435a (1), 720 (1), 746 (1), 755 (1),
7965 (1), 3070A (1), 3478 (1), BS41748{\), BS77711 (4), BS77570 (4); Ellis, J.L. 20452
{3)\ Elmer A.D.E. 5575(1), 9695(1), 75476(1), 75626(1), 75577(1), 77557 (1), 75255

(1) , 20569 (16); Endert, FH. 1762 (16), 5143 (16); Ernst, A. 1118 (16); Escritor, L.
5527452 (1); Esquirol, J. 711 (2); Etuge, M. 1973 (20), 2057 (20); Eyma, P.J. 1826 (7),
2574 (7).

Faden, R.B. & Faden, A.J. 76/494 (3); Fenix, E. BS26049 (3), BS28106 (3); Feuilletau de
Bruyn, W.K.H. 335 (1); Forrest, G. 15699 (2); Fosberg, FR. 57278 (3); Frodin, D.G. &
Bin Ismawi, O. 2077 (16).

Gachalian, F. 147 (1); Gafui, I.H. & collectors BSIP14759 (14), BSIP14763 (14); Gallatly,
G. 1035 (16); Gardner, G. 606 (3); Geesink, R. cS: Phengklai, C. 6082 (16), 6257 (16);
Geqffray, M. 95 (3); Grey-Wilson, C. & Grey-Wilson, C.M. 3016 (3); Grupe, D.A. 200
(3); Guinea, E. 1408 (20).

Haegens, R.M.A.P. & Klazenga, N. 515 {\5y, Hallier, H.G. 91 (16), 4725 {\);Flamid H18 (16);
Hansen, C. 17 (15); Hardial, S. & Sidek 474 (16); Harris, D. 5815 (20); Haviland,
G.D. s.n. (15); Haviland, G.D. & Hose, C. 3525 R (15); Henderson, M.R. 19406 (10),
20277 (16), 22552 (10), 22580 (10), 22875 (16), 25010 (10), 25205 (16), 29079 (16);
Hennipman, E. 610g (7); Henriques, J.A. 5 (20), 12 (20); Heniy, A. 1916 (3), 12280

(2) , 12705 (2), 75497 (2); Hepper, FN. & Maley, J. 8073 (20); Hicks, D. 151 (1);
Hochreutiner B.P.G. 7976 (16); Holttum, R.E. 15125 (16); Hooker, J.D. & Thomson,
T. s.n. (2), 2161 (2); Hoover, W.S. et al. HW11417 (5), Deden 757 (5); Horsfield, T.
Cyrtandra 3 (5), 55 (5); Hotta, M. et al. 313 (18), 579 (18), 524 (1 8), 559 (1 8), 402 (18),
659 (18); Hotta, M. 14402 (15), 75257 (15); Hotta, M. & Okada, H. 247 (16), 576 (18),
1087 (18); Hsieh, C.F et al. s.n. (3).

Iboet 221 (6); Idjan 175 (1); Idzumi. H. & Togashi, M. s.n. (16); Ilias bin Paie S. 28047 (16),
5.25657 (16); Imaichi, R. etal. 2 (16), 30 (15), 55 (15); Imin, K. FR163208 (11).

Jaag, O. 1600 (19); Jacques-Felix, H. s.n. (20); Jacobson, E. 199 (5), 275 (16); Jaeger, P. 227
(20); Jaheri 1522 (15); Jamree et al. S. 82239 (1 6); Jayasuriya, A.H.M. 843 {3);Jemree,
5. & Enjah, A. S. 85584 (16); Jemree, S. et al. S. 82056 (16), S. 84036 (16); Jeswiet, J.
670 (5); Julaihi S. 86868 (15); Julaihi et al. S. 76938 (15); Julia, 5. et al. S. 93376 (15),
5.94757 (15), 5.99565 (15); Junghulm, F.W. 3 (16).

Revision of Epithema

227

Kamarul Hisham, M. FRI67230 (16); Kanehira, R. & Hatusima, S. 11848 (1), 13143 (1);
Kasem 641 (3); Kato, M. et al. C.5728 (7), C.5232 (7), C.5144 (7), C.6007 (7), C.6908
(1), C.11441 (7), BJ1733 (16); Keenan, J. et aL 798 (16); Kenfack, D. 259 (20); Kerr,
A.F.G. 2655 (16), 6381 (16), 7657(16), 12385 (16), 13053 (3), 13183 (16), 15775 (10),
15893 (10), 19385 (16); Kessler 226 (16); Kie\v, R. et al. RK4321 (4); Kiew, R. RK525

(15) , RK1231 (10), RK1347 (10), RK1349 (16), RK1423 (16), RK1498 (11), RK1566

(16) , RK1570 (16), RK1584 (10), RK1590A (10), RK2145 (16), RK2287 (16), RK2369
(16), RK3653 (16), RK3712 (16), RK4711 (16), RK4759 (4), RK5040 (16), RK512()
(16X RK5254 (10), RK5276 (16); Kiew, R. & Anthonysamy, S. RK3020 (16); Kiexv, R.
& Lim, S.F. RK4138 (4), RK4211 (16); King, G. s.n. (2); King’s Collector 5872 (16),
7046 (16); Kjellberg, G.K 195 (16), 314 (6), 315 (6), 1299 (16), 2616 (1), 2618 (19);
Klaine, R.P. 1517 (20), 2666 (20); Knedl, M. 9/365 (20), 721 (20); Kochummen, K.M.
FRI26221 (11); Kokawa, S. 6381 (16); Kokawa, S. & Hotta, M. 551 (16), 557 (16), 565
(16), 1073 (4), 1125 (4); Koorders, S.H. 29801 (16), 40438 (16); Kooy, C.W. 403 (5),
789 (5); Kornassi 482 (1), 621 (7), 967 (7); Korthals, P.W s.n. (18), 692 (18); Koster,
C. BW10773 {\\ BW13665 (1); Kostemians, AJ.G.H. 1103 (16), 7546(16), 1401 (16),
18053B (5), 18629 (5), 18839 (5), 21338 (16); Kostermans, A.J.G.H. & Soegeng 914
(1); Kulkarni, B.G. 108673 (13); Kunstler, H. 983 (16); Kuswata 105 (5), 121 (5), 213
(5), 775 (16).

Lace, J.H. 4892 (2); Lai & Rantai et al. S. 66009 (16); Lamb, A.L. 8/85 (16); Larsen, K. et al.
722 (16), 2923 (3), 3363 (16), 30967 (16), 41243 (16), 41413 (16), 41450 (16), 41713
(10), 42245 (10), 42312 (10), 42471 (10), 42579 (10), 42584 (10), 43589 (3), 43771
(3), 44271 (10), 46055 (16), 46161 (2), 46426 (3), 46861 (3); Le Testu, M.G. 5449 (20),
7018 (20); Lehmann, PF. S.30091 (15), PFL364 (16), PFL596/S, 30372 (15), S.30091

(15) ; Leong, P. et al. PL215 (15), PL320 (15); Lesger 359 (16); Letouzey R. 7722 (20),
8732 (20), 9238 (20), 13198 (20), 13973 (20), 15020 (20), 15026 (20); Lim, S.P et al.
580 (16), LSP675 (4), LSP681 (4), LSP738 (16); Lobb, T. 387 (3); Loeters, J.J. 1625 (5);
Loher, A, 1554 (3), 1555 (3), 7556 (3), 4231 (I ), 6665 (3), 6669 (3); Lopez, G. BS42031
(1), BS42049 (1), BS41343 (9); Lorzing, J.A. 17359 (16); Lucas 1025 (3); Lutjeharms,
W.J. 4598 (15), 5112 (15); Lye, KA. & Katende, A.B. LYE22885 (20).

Mackinder, B.A. 303 (20); Madulid, D.A. et al. 11563 (9); Maier Sarip, R. 162 (5); Mann, G.
2345 (20); Maradji 482 (16); Marcan, A. 1869 (3); Ma’roef, A. AM. 250 (16); Martin,
PJ. S.39273 (15); Maxwell, J.F s.n. (16), 72-362 (16), 73-107 (3), 73-285 (16), 74-814

(16) , 75-958 (1 6), 84-55 (1 6), 86-723 ( 1 6), 86-893 (10), 87-1154 (3), 89-920 (2), 93-886
(16), 95-536 (3), 99-278 (2); McGregor, R.C. BS19951 (1), BS43566 (3); McKee, H.S.
1958 (3), 6577 (3); Meekiong, K. SBC778 (1 6); Meijer; W. 5657 (1 8), 9133 (19), 10045
(1), SAN20756 (16); Mendoza, D.R, & Convocar, P P. 226 (12), 297 (12); Mendum,
M. et al. 34 (5), 45 (1), 775 (1 ), 180 (1 ); Merrill, E.D. 52 (3), 4216 (3), 5106 (1 ), 8222
(1); Middleton, D.J. et al. 4055 (16), 4284 (3), 4330 (16), 4545 (16), 4428 (16), 4544
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Nadiah, 1. et al. SAN149136 (4); Nair, K.K.N. 1354 (3); Namba, T. & Mikage, M. 82-413

228

Card. Bull. Singapore 67(1) 2015

(3); Ncmcikorn, W 4240 (3); Nanakorn, W et al. 4664 (2); Narayanaswami, M.A. 478

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Revision of Epithema

229

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Gardens’ Bulletin Singapore 67(1): 231-250. 2015
doi: 10.3850/S2382581215000186

231

Synoptic overview of Acacia sensu lato (Leguminosae:
Mimosoideae) in East and Southeast Asia

B.R. Maslin

Western Australian Herbarium, Department of Parks and Wildlife,
Locked Bag 104, Bentley Deliveiy Centre, Western Australia 6983

Baice.IVIaslin@dpaw.wa.gov.au
Honorary Research Associate

Herbarium, Singapore Botanic Gardens, National Parks Board,

1 Cluny Road, Singapore 259569

ABSTRACT. Recent research shows that the formerly broadly circumscribed, pantropical genus
Acacia Mill. (Leguminosae: Mimosoideae) is polyphyletic and should be treated as comprising
at least five genera, namely. Acacia Mill, sensu stricto, Acaciella Britton & Rose, Mariosousa
Seigler & Ebinger, Senegalia Raf. and Vachellia Wight & Arn. The indigenous flora of Acacia
sensu lato in East and Southeast Asia comprise 52 species: 32 species (38 taxa) of Senegalia, 12
species of Acacia sensu stricto and eight species of Vachellia. These species are listed and their
geographic distributions given, showing that Acacia sensu lato is unevenly distributed across
the region, with centres of species-richness in Indonesia, Myanmar and Thailand. A summary
of the classification histoiy of Acacia sensu lato is provided and nomenclatural impacts of the
recent retypification of Acacia briefly discussed.

Keywords. Classification, nomenclature, phylogeny, Senegalia, Vachellia

Introduction

Molecular and other evidence has shown that the fonnerly broadly circumscribed,
pantropical genus Acacia Mill. (Leguminosae: Mimosoideae) is polyphyletic and
should be treated as comprising at least five genera, namely, Acacia Mill, sensu stricto,
Acaciella Britton & Rose, Mariosousa Seigler & Ebinger, Senegalia Raf. and Vachellia
Wight & Arn. (Miller & Seigler, 2012). Collectively, these five genera are called
Acacia sensu lato in the discussion below. The name Acacia is now conserved with a
new type (McNeill & Turland, 2011), an action which has had global nomenclatural
repercussions. AJthough increasingly understood and adopted, the new classification
for Acacia sensu lato has not yet been universally adopted, despite the fact that names
congment with the new classification are now available for most currently accepted
taxa.

The indigenous species of Acacia sensu lato in East and Southeast Asia comprise
52 species, mostly belonging to Senegalia (see Appendix 1), These species are unevenly
distributed across the region with centres of species-richness in Indonesia, Myanmar
and Thailand (see Appendix 2). The tenu East Asia here refers to China and Taiwan
where species of Acacia sensu lato occur both naturally and as introductions. The term

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Card. Bull. Singapore 67(1) 2015

Southeast Asia includes the countries from Papua New Guinea through Indonesia,
Philippines, Brunei, Vietnam, Laos, Cambodia (Kampuchea), Singapore, Malaysia,
Thailand to Myanmar (Burma). Although New Guinea is geographically part of
Australasia, it is included here for convenience and because the island is included in
the largest Flora project in Southeast Asia, namely. Flora Malesiana. The species of
Acacia sensu lato of this whole region are currently under review (Maslin, in prep.),
a work that benefits greatly from the excellent taxonomic foundation provided by the
late Ivan Nielsen for Southeast Asian taxa (Nielsen, 1980, 1981, 1985a, 1985b, 1992)
and the treatments by Sun & Chen (1990) and Wu & Nielsen (2010) for taxa from
China.

The aims of this communication are to summarise the rationale underlying the
new classification and updated nomenclature of Acacia sensu lato and to show how
this structure and these names apply to the indigenous species of East and Southeast
Asia. This will provide an introduction to the above mentioned taxonomic review
and will facilitate implementation of the new classification and nomenclature for the
region.

Phylogeny and classification of Acacia sensu lato

As defined until relatively recently, the genus Acacia comprised a very large group of
about 1350 species distiHbuted throughout tropical and warm temperate areas of the
world, occurring on all continents except Antarctica (web. ref 1). However, during
the past decade the genus has undergone fragmentation, a process that has been driven
largely by evidence derived from molecular phylogenetic studies.

According to Bentham (1840), during the 80 years following its original
description by Miller (1754) Acacia had become an “unwieldy, ill-defined, and
comparatively unnatural assemblage of plants”. Bentham (1842) remedied this situation
by restricting the name Acacia (today’s Acacia sensu lato) to Mimosoid plants having
indefinite, free stamens, a definition that has persisted to modem times. In a series of
subsequent publications, culminating in his 1 875 magnum opus, Bentham did much to
clarify not only the definition of Acacia but also its internal classification (Bentham,
1875). During the ensuing 60 years 15 new genera were described but these were
ultimately treated as congeneric with Acacia sensu lato. A discussion of this generic
history is outlined in Maslin et al. (web ref. 1).

In 1986 Pedley published a reassessment of the classification of Acacia,
dividing the genus into three, namely. Acacia sensu stricto, Senegalia diwd Racosperma
(DC.) Mart. These genera corresponded to three subgenera of Acacia sensu lato that
had previously been recognised by Vassal (1972), namely. Acacia subgenera Acacia,
Aculeiferum Vassal and Heterophyllum Vassal (=subgen. Phyllodineae (DC.) Seringe)
respectively. At the time, Pedley ’s classification was not widely adopted for reasons
that are outlined in Maslin (1989) and Maslin et al. (web ref. 1). One of these reasons
was that the evidence presented was considered inconclusive and/or incomplete.

Synoptic overview oi Acacia sensu lato

233

especially considering the significant impact to global nomenclatural that would flow
from such a division.

However, since 1986 a number of broad-based, comparative studies (particularly
using chloroplast and nuclear DNA) provided evidence that enabled more robust
and infoinied decisions to be made concerning the phytogeny and classification of
Acacia (see publications listed in Miller & Seigler, 2012). These studies included not
only species of Acacia sensu lato and Faidherbia A.Chev. (which together comprise
tribe Acacieae) but importantly, also included representatives from the related tribes
Ingeae and Mimoseae. As summarised in Miller & Seigler (2012) this genetic data
has consistently demonstrated that Acacia sensu lato is polyphyletic and comprises at
least five monophyletic groups, which each warrant recognition as a distinct genus.
These genera are shown in Table 1 and comprise (1) two small New World endemic
genera, Acaciella (resurrected by Rico Arce & Bachman, 2006) and Mariosousa
(new genus described in Seigler et al., 2006), (2) two relatively large pan-tropical
genera, Senegalia (resun-ected by Pedley, 1986) and Vachellia (which contains the
original type species of Acacia, A. nilotica (L.) Willd. ex Del.), and (3) the enormous,
predominantly Australian genus, Acacia sensu stricto (syn. Racosperma). Vachellia
is taxonomically well-removed from the other genera of Acacia sensu lato, being
nested within a paraphyletic tribe Mimoseae; as noted by Miller & Seigler (2012)
there are two, well-supported subclades within Vachellia, one of African species and
the other American, to which the type of Vachellia, V farnesiana (L.) Wight & Am.,
belongs. The other four genera are in a paraphyletic grade with genera of tribe Ingeae
(fide Miller & Seigler, 2012). It is this five-genus classification fox Acacia sensu lato
that is generally accepted today. In addition to the phylogenetic evidence based on
nucleotide data, a range of morphological, biochemical, palynological and other data
help support, define and characterise the five genera (see Pedley, 1986; Chappill &
Maslin, 1995; Maslin et al., 2003).

Notwithstanding the above it is possible that further generic segregates will
be recognised within uicacia sensu lato, particularly as more taxa are added to the
genetic datasets. Miller & Seigler (2012) have already suggested a possible new
genus in South America, segregated fi*om Senegalia, to accommodate species allied
to S. skleroxyla (Tussac) Seigler & Ebinger. While major generic realignments are not
anticipated for East and Southeast Asia (but see note below concerning Delaportea
Gagnep.), it is noted that scarcely any species from the region has been included in
existing comparative genetic studies of Acacia sensu lato

Nomenclature of Acacia sensu lato

In July 2011 the Nomenclature Section of the XVII International Botanical Congress
(IBC) in Melbourne, Australia, voted with a clear majority (68%) to accept the Vienna
Code (McNeill et al., 2007) that lists Acacia Mill, with a conserved type (McNeill &
Turland, 2011). This action ended a long debate that began following the publication

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Card. Bull. Singapore 67(1) 2015

of a formal proposal by Orchard & Maslin (2003) to replace the original type of
Acacia, the African species A. scorpioides (L.) W.F. Wight (=A. nilotica), with a new
type, the Australian species, A. penninei^is Sieber ex DC. A major consequence of the
Melbourne IBC decision is that the ndmQ Acacia now applies to the large, predominantly
Australian group that was formerly called Acacia subgen, Phyllodineae; this group is
referred to here as Acacia sens. str. and the name Racosperma is a synonym of it.
Additionally, the name Vachellia is the correct name for the smaller, pan-tropical group
that was fonnerly called Acacia subgen. Acacia. Further infoiination concerning this
nomenclatural issue is provided on the Worldwide Wattle website (web ref 2).

Combinations for almost all the currently recognised species of Acacia sensu
lato now referable to Acaciella, Mariosousa, Senegal ia and Vachellia have been made,
with only 13 species (10 from Madagascar and three from Africa) outstanding (web
ref 3). The WorldWideWattle website (web ref 4) provides a list of names under
the new classification, and where appropriate, each name is cross-referenced to its
analogue in Acacia when that group was treated as a single genus.

After the Vienna IBC, the ‘new’ generic names replacing Acacia sensu lato had
begun to appear in a range of publications and web databases, and the trend continues
today; however, there seems to have been a lesser uptake of these names by herbaria. As
summarised by Maslin (201 1 ), even prior to the Melbourne IBC the name Vachellia had
been adopted, in lieu of Acacia, in many publications including Flora treatments, field
guides, scientific research papers and books. One particularly important publication
to adopt the new nomenclature early was Mabberley s Plant-book 3^^* ed. (Mabberley,
2008), which is a primaiy reference source for the correct names of vascular plant
genera and families of the world. The new nomenclature has now been adopted in
some large web databases such as the National Center for Biotechnology Infomiation
(web ref. 5) and will be incorporated into the Catalogue of Life database and available
online by mid 2015 (Y. Roskov, pers. comm.). Others web resources such as Tropicos
(web ref 6) and The Plant List (web ref 7) currently adopt a half-way approach by
listing many names as accepted under both Acacia and Vachellia/Senegalia. Some
important legume-centric databases on the web still maintain the old nomenclature,
e.g. the online version of Legumes of the World (web ref 8) and the International
Legume Database & Information Service (ILDIS) (web ref. 9). Furthennore, the
online version of Index Nominum Genericorum (web ref. 10) still lists A. scorpioides
as the type of Acacia. There are of course many possible reasons why the uptake of the
new generic nomenclature has been slow, erratic or has not occurred at all. Resource
and time limitations are two of the more obvious reasons, but some databases may
simply not have the capacity to be changed (ILDIS is presently in this latter category).

In East and Southeast Asia the new names have begun to appear on the web
with Senegalia being listed for both the Philippines (web ref. 11) and for the Hengduan
Mountains region of south-central China (web ref 12). Also, the new generic
nomenclature has been adopted in the checklist of plants of China that is expected
to be soon published and in the checklist of the plants of India that is cun'ently in
preparation and which is expected to appear online during 2015, and published in
hard copy about a year thereafter. Australia has adopted the new nomenclature in its

Synoptic overview oi Acacia sensu lato

235

national names database (web ref. 13) and Australian herbaria have by and large done
likewise with their specimen records (web ref. 14). However, this was a relatively
easy achievement for Australia because so few name and specimen records had to be
changed, with just 1 1 indigenous species being affected (see Table 1).

It is recognised that it often takes a long time for taxonomic and nomenclatural
changes to be accepted and implemented. For herbaria the adoption of such changes
can be especially troublesome not only because of the large amount of work and
resources needed to redetermine specimens and (often) modify database records,
but also because of the disruption caused by having to re-aiTange specimen storage
systems. In the case of Acacia sensu lato these problems are compounded due to
its very many species and global distribution (thus many herbaria are aftected), and
also because of the drawn-out debate over the application of the name Acacia itself
While some people may still wish that change had not occun'ed (a sentiment that
is understandable) the fact remains that the international botanical nomenclatural
community has voted in favour of Acacia retypification. It is therefore appropriate that
the new nomenclature for Acacia sensu lato be adopted universally so that stability can
be provided for this unportant group of legumes. There is now a particular imperative
to achieve this nomenclatural stability because the global legume community is
discussing ways to produce a new phylogenetic classification for the Leguminosae,
one that will incorporate morphological data hito the phytogeny (Bruneau et af, 2013).
One of the basic starting points for this project will be the development of a core list of
taxa; Acacia sensu lato, whose species constitute about a third of the total number for
subfamily Mimosoideae, will be a significant component of that list.

The indigenous species of Acacia sensu lato in East and Southeast Asia

Current evidence shows Acacia sensu lato to be represented in the East and Southeast
Asia by 52 native species accoimiiodated in three genera, namely. Acacia sens,
str., Senegalia and Vachellia, The species are listed, together with their geographic
distributions, in Appendices 1 and 2. However, it should be noted that some modification
to these data can be expected to occur within the context of the review of Acacia sensu
lato for East and Southeast that is currently in progress (Maslin, in prep.).

Acacia sensu stricto

Acacia sensu stricto (1073 species in total) is largely restricted to Australia where
1063 species occur (Table 1), making it the largest genus of vascular plants on that
continent. Only 19 taxa (representing 18 species) occur naturally outside Australia
(Brown et al, 2012), 12 in Asia and seven on islands of the Pacific.

All 12 Asian species are geographically restricted, one occurring in East Asia
{Acacia confusa Merr. from Taiwan and the Philippines) and 1 1 in Southeast Asia.
There are four endemic taxa, three in the south of the region, where Acacia sp. (Wetar)

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Card. Bull. Singapore 67(1) 2015

and A. wetarensis Pedley are restricted to the small Indonesian island of Wetar and A.
peregrin alls M.W. McDonald & Maslin which is confined to New Guinea, and one in
the northeast (the above mentioned A. confusa). All eight species that extend beyond
the region occur also in Australia. There are no indigenous species of Acacia sensu
stricto in mainland China but a few are cultivated there (fide Wu & Nielsen, 2010).

The indigenous species of Acacia sensu stricto in East and Southeast Asia are
shrubs or trees with phyllodinous foliage (bipinnate foliage occurs on a few introduced
species in Asia). Like species of Vachellia they are generally found in drier habitats
than those of Senegalia.

Some taxa of this genus foiin a significant component of the plantation forestry
industry in parts of Southeast Asia, most notably in Indonesia and Vietnam, where
Acacia crassicarpa A.Cuim. ex Benth., A. mangium Willd. and A. auriculiformis
A.Cunn. ex Benth. x mangium are the main ones grown for pulp and solid wood
products (see Griffin et al., 2011 for overview).

Senegalia

Senegalia (201 species in total) has a pantropical distribution and is represented by 45
species in Asia as a whole (19 in India), 68 in Africa, 102 in the Americas and 2 taxa
in Australia (see Table 1 ).

There are 38 indigenous taxa of Senegalia (representing 32 species) in East
and Southeast Asia, making it by far the largest group of Acacia sensu lato within the
region. Thirteen taxa occur in East Asia with four, Senegalia delevayi (Franch.) Maslin
et al. (van delavayi and van kiinmingensis (C.Chen & H.Sun) Maslin et al.), S. teniana
(Harms) Maslin et al. and S. yunnanensis (Franch.) Maslin et al., endemic in China.
Thirty three taxa occur in Southeast Asia with almost half of them endemic to the sub-
region, namely, Senegalia borneensis (I.C.Nielsen) Maslin et al., S. comosa (Gagnep.)
Maslin et al., S. donnaiensis (Gagnep.) Maslin et al., S. kekapur (I.C.Nielsen) Maslin
et al., S. kostermansii (I.C.Nielsen) Maslin et al., S, meeboldii (Craib) Maslin et al.,
S. merrillii (I.C.Nielsen) Maslin et al., S. pala\\>anensis (I.C.Nielsen) Maslin et al.,
S. pluricapitata (Steud. ex Benth.) Maslin et al., S. pluriglandulosa (Verde.) Maslin
et al., S. pseiidointsia (Miq.) Maslin et al., S. sulitii (I.C.Nielsen) Maslin et al., S.
tawitaMdensis (I.C.Nielsen) Maslin et al., S. thailandica (I.C.Nielsen) Maslin et al. and
S, verheijenii (I.C.Nielsen) Maslin et al. These endemic taxa are scattered generally
tliroughout the sub-region, occurring in all countries except Brunei (which has no
indigenous Acacia sensu lato recorded). Indonesia has the highest concentration of sub-
regional endemics with nine species recorded, but only Senegalia kostermansii and S.
verheijensii are restricted to that country. Almost all of the 16 species of Senegalia that
extend beyond the region range westward to India and nearby countries, with 10 also
ranging north to China. Only one taxon, Senegalia pennata subsp. kerrii (I.C.Nielsen)
Maslin et al., extends to Australia, which has a very poor representation of indigenous
species of Senegalia (just two taxa, the endemic S, albizioides (Pedley) Pedley and S,
pennata subsp. kerrii).

Synoptic overview oi Acacia sensu lato

237

Species of Senegalia are characterised by having bipinnate leaves and cauline
prickles. Most of the indigenous species of East and Southeast Asian are woody
lianes with only the widespread Senegalia catechu (L.f.) P.J.H.Hurter & Mabb., S.
chundra (Roxb. ex Rottler) Maslin and S. ferruginea (DC. ) Pedley (all of which have
prickles at the nodes) and the geographically restricted, endemic S. kostermansii
from Indonesia and S. teniana from China (both of which have scattered prickles)
seemingly obligate shrubs or trees which lack scandent branches. Most species have
broad, flat, ±chartaceous and straight pods but in Senegalia thailandica the pods
are slightly inflated and tightly curled. These species can be arranged in four sub-
groups (defined principally by prickle distinbution and leaflet characteristics) centred
on Senegalia pennata, S. caesia, S. andamanica (l.C.Nielsen) Maslin et al and S.
catechu respectively. The one species with very different caipological features is
the widespread and variable Senegalia rugata (Lam.) Britton & Rose (syn. Acacia
concinna (Willd.) DC.). In this species the pods are smooth, thick and fleshy when
fresh but they dry characteristically wrinkled, blackish and with a very hard texture.
This species name has about 10 heterotypic synonyms and fiirther study is needed to
reassess the taxonomic status of Senegalia rugata. Senegalia albizioides (restricted to
Australia) is currently the only recognised close relative of S. rugata.

Vachellia

Vachellia (163 species in total) has a pantropical distribution and is represented by
30 species in Asia as a whole (11 in India), 84 in Africa, 57 in the Americas and 8 in
Australia (see Table 1).

There are just eight indigenous species of Vachellia in Southeast Asia (none
occurs in East Asia), making it the least well-represented of the three genera within
the region. There are five endemic species, all of which occur m northern areas, three
restiicted to Myanmar {Vachellia inopinata (Prain) Maslin et al., V. kingii (Prain)
Maslin et al. and V. myaingii (Lace) Maslin et al.: all these species are poorly known
and require study to reassess their taxonomic status), one to Thailand {V siamensis
(Craib) Maslin et al.) and one widespread {V. harmandiana (Pierre) Maslin et al.
which occurs in Cambodia, Laos, Thailand, Vietnam). The three species that extend
beyond the region, namely, Vachellia nilotica subsp. indica (Benth.) Kyal. & Boatwr.,
V leucophloea (Roxb.) Maslin et al. and V tomentosa (Rottler) Maslin et al., range
westward to India with V. nilotica subsp. indica extending to the Middle East.
Excluding Vachellia harmandiana and V leucophloea, the only indigenous species of
Vachellia that is widespread in Southeast Asia is V. tomentosa.

Gagnepain (1911, 1952) and Craib (1927) described species under new generic
names, Delaportea and Nimiria Prain ex Craib respectively. At present these species
are treated as conspecific with Vachellia harmandiana, V. leucophloea and V. siamensis.
They appear to fonn a natural group that is recognised by an unusual combination of
characters, namely, stipules spinescent (characteristic of Vachellia) and heads arranged
in open, terminal panicles (characteristic of many species of Senegalia). Further field,

238

Card. Bull. Singapore 67(1) 2015

morphological and genetic study is needed to reassess the taxonomic status of this
group and to determine if it is appropriate to resurrect the genus Delaportea (with
Nimiria as a synonym),

Vachellia farnesiana (L.) Wight & Am. has been recorded as an introduction
(sometimes naturalised) in a number of countries of East and Southeast Asia (see
Nielsen, 1981, 1985b, 1992; Wu & Nielsen, 2010). It is a native of tropical America
and comprises three varieties (Seigler & Ebinger, 2005). Although it is not known
what varieties occur in Asia it is most likely the typical one (D. Seigler, pers. comm.).

The indigenous species of Vachellia in Southeast Asia are shrubs or trees
characterised by having bipinnate foliage subtended by spiny stipules. Like species of
Acacia sens. str. they are generally found in drier sites than those of Senegalia.

ACKNOWLEDGEMENTS: Prof Phan Ke Loc (Vietnam National Herbarium, Hanoi) is
thanked for discussions that provided the motivation for me to prepare this communication.
The following colleagues are thanked for providing constructive comments on the manuscript:
Prof David Mabberley (Sydney), Dr Kevin Thiele (W.A. Herbarium, Perth) and Dr Boon-
Chuan Ho (Singapore Botanic Gardens). The following herbaria are thanked for allowing
me access to their collections: Herbarium Bogoriense, Cibinong, Indonesia (BO); Bangkok
Herbarium, Chatuchak, Thailand (BK); The Forest Herbarium Bangkok, Chatuchak, Thailand
(BKF ); Chiang Mai University Herbarium, Thailand (CMU); Hanoi Herbarium, National Center
for Natural Sciences and Technology, Hanoi, Vietnam (HN); Vietnam National Herbarium,
Hanoi University, Hanoi, Vietnam (HNU); Forest Research Institute Malaysia Herbarium,
Kuala Lumpur, Malaysia (KEP); Kunming Institute of Botany, Chinese Academy of Sciences
Herbarium, Kumning, China (KUN); Singapore Botanic Gardens Herbarium, Singapore
(SING); Taiwan Forestry Institute, Taipei (TAIF); Institute of Tropical Biology Herbarium, Ho
Chi Minh City, Vietnam (VNM).

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241

2. Acacia Name Issue. http://www.worldwidewattle.com/infogalleiy/nameissLie/index.php.

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Appendix 1. Indigenous taxa of Acacia sensu lato currently recognised for East and Southeast
Asia.

See Introduction for definition of the terms East and Southeast Asia. Only selected synonyms
are given in this list.

ACACIA Mill., Gard. Diet. Abr., ed. 4, [25] (1754), nom. cons. TYPE: Acacia penninervis
Sieber ex DC. {typ. cons., vide Regnum Veg. 146: App. Ill, 286. 2006).

Racosperma (DC.) Mart., Hort. Reg. Monac. Semin. 4 (1835). LECTOTYPE: Racosperma
pennineiA^e (Sieber ex DC.) Pedley, Bot. J. Linn. Soc. 92: 239 (1986). Based on Acacia sect.
Phyllodineae DC., Prodr. 2: 448 (1825).

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Card. Bull. Singapore 67(1) 2015

Acacia aur iculif or mis A.Cmm. ex Benth., London J. Bot. 1: 377 (1842).

Distribution. Southeast Asia: Indonesia (Moluccas), Papua New Guinea. Australia: Northern
Territory & Queensland.

Introduced/cultivated/naturalised. Americas (Florida, Panama), Africa, Bangladesh, Brunei,
China, India (inch Andaman Is.), Indonesia, Laos, Malaysia, Myanmar, Nepal, Pakistan,
Philippines, Singapore, Sri Lanka, Vietnam.

Acacia confusa Men*., J. Sci. (Bot.) 5: 27 (1910).

Distribution. East Asia: Taiwan. Southeast Asia: Philippines.

Cultivated/introduced. China, Hawaiian Islands, India, Indonesia, Japan (southern Ryuk5m
Islands), Mauritius, Malaysia, Northern Marianas, Seychelles, Vietnam.

Acacia crassicarpa A.Cunn. ex Benth., London J. Bot. 1: 379 (1842).

Distribution. Southeast Asia: Papua New Guinea. Australia: Queensland.

Cultivated. Australia (Northern Territory), Vietnam.

Acacia leptocarpa A.Cunn. ex Benth., London J. Bot. 1: 376 (1842).

Distribution. Southeast Asia: Papua New Guinea. Australia: Northern Territory, Queensland,
Western Australia.

Acacia mangium Willd., Sp. PL, ed. 4, 4(2): 1053 (1806).

Distribution. Southeast Asia: Indonesia (New Guinea, Moluccas), Papua New Guinea.
Australia: Queensland.

Introduced/naturaUsed. Americas (Nicaragua), Australia (Northern Temtory), Bangladesh,
Brunei, China, Central America (Nicaragua), hidia, Philippines, Singapore, Vietnam.

Acacia oraria F.Muell., Fragm. 11: 66 (1 879).

Distribution. Southeast Asia: Indonesia (Flores, Timor). Australia: Queensland.

Acacia peregrinalis M.W.McDonald & Maslin, Nuytsia 14(3): 455 (2002).

Distribution. Southeast Asia: Indonesia (New Guinea), Papua New Guinea.

Acacia pubirhachisVQdlQy, Contr. Queensland Herb. 15: 15 (1974).

Distribution. Southeast Asia: Papua New Guinea. Australia: Queensland.

Acacia simsii A.Cunn. ex Benth., London J. Bot. 1: 368 (1842).

Distribution. Southeast Asia: Indonesia (New Guinea), Papua New Guinea. Australia:
Northern Territoiy, Queensland.

Acacia spirorbis subsp. solandri (Benth.) Pedley, Austrobaileya 3: 216 (1990).

Distribution. Southeast Asia: Papua New Guinea. Australia: Queensland.

Acacia wetarensis ?Qd\Qy, Contr. Queensland Herb. 18: 18 (1975).

Distribution. Southeast Asia: Indonesia (Wetar).

Acacia sp. (Wetar).

Distribution. Southeast Asia: Indonesia (Wetar).

Note. This entity is noted by Nielsen (1992: 59) [inder Acacia leptocarpa. It appears close to
the glabrous variant of A. elachantha M.W.McDonald & Maslin (McDonald & Maslin, 1997),
and preliminary studies suggest that it may represent a distinct taxon.

Synoptic overview oi Acacia sensu lato

243

SENEGALIA Raf., Sylva Tell. 1 19 (1838). LECTOTYPE: Mimosa Senegal L. {vide Britton &
Rose, N. Amer. FI. 23: 106. 1928) {Senegalia Senegal (L.) Britton [Senegalia triacantha Raf.,
nom. illeg.])

Senegalia andamanica (l.C.Nielsen) Maslin, Seigler & Ebinger, Blumea 58: 40 (2013). -
Acacia andamanica l.C.Nielsen, Adansonia ser 2, 19: 354 (1980).

Distribution. Southeast Asia: Thailand. South Asia: Andaman Islands, Nicobar Islands.

Senegalia borneensis (l.C.Nielsen) Maslin, Seigler & Ebinger, Blumea 58: 40 (2013). -Acacia
borneensis l.C.Nielsen, Opera Bot. 81: 20, fig. 9 (7-13) (1985).

Distribution. Southeast Asia: Indonesia (Kalimantan), Malaysia (Sabah).

Senegalia caesia (L.) Maslin, Seigler & Ebinger, Blumea 58: 40 (2013). - Mimosa caesia E.,
Sp. PI. 1: 522 (1753). -Acacia caesia (L.) Willd., Sp. PI., ed. 4, 4(2): 1090 (1806).
Distribution. East Asia: China (Guangdong, Hainan, Sichuan, Yunnan), Taiwan. Southeast
Asia: Cambodia, Laos, Myamnar, Thailand, Vietnam. South Asia: Bangladesh, Bhutan, India,
Sri Lanka.

Note. Although Acacia caesia van subnuda (Craib) l.C.Nielsen is recognised by Nielsen
(1981 : 53) and Sanjappa (1 992: 37), it is regarded as conspecific with .^4. caesia by Chakrabarty
& Gangopadhyay (1996: 604) and Wu & Nielsen (2010: 57). Maslin et al. (2013) did not
recognise the variety in Senegalia. Further study is required to reassess the taxonomic status
of this variety.

Senegalia catechu (L.f ) P.J.H.Hurter & Mabb. in Mabberley’s Plant-book ed. 3, 1021 (2008).
-Mimosa catechu L.f, Suppl. PI. 439 (1782 [1781 publ. Apr. 1782]). - Acacia catechu (L.f.)
Willd., Sp. PI., ed. 4, 4(2): 1079 (1806).

Distribution. East Asia: China (Yunnan). Southeast Asia: Myanmar, Thailand (uncertain, fide
Lock & Heald 1994: 33). South Asia: Bangladesh, Bhutan, India, Nepal, Pakistan, Sri Lanka.
Cultivated/introduced. China (Fujian, Guangdong, Guangxi, Hainan, Yunnan, Zhejiang),
Indonesia (Java), Japan (southern Ryukyu Islands), Mauritius, Philippines, Taiwan, Vietnam.

Senegalia chiindra (Roxb. ex Rottler) Maslin, Nuytsia 22(6): 466 (2012). - Mimosa chundra
Roxb. ex Rottler, Neue Schriften Ges. Naturf. Freunde Berlin 4: 207 (1803). - Acacia chundra
(Roxb. ex Rottler) Willd., Sp. PI. ed. 4, 4(2): 1078 (1806).

Acacia sundra (Roxb.) DC., Prodr. 2: 458 (1825). - Mimosa siindra Roxb., PI. Corom. 3: 19,
tab. 225 (1811).

Distribution. Southeast Asia: Myamnar. South Asia: India, Sri Lanka.

Introduced/naturalised. Australia: Northern Territory, Reunion Island.

Note. Despite their similar-sounding epithets. Mimosa chundra andM sundra were independent
descriptions based on different types,2i4fe Maslin et al. (2013: 466). The complex nomenclatural
history involving these names is discussed by Kshirsagar (web ref. 15).

Senegalia comosa (Gagnep.) Maslin, Seigler & Ebinger, Blumea 58: 40 (2013). - Acacia
comosa Gagnep., Notul. Syst. (Paris) 2: 113 (1911).

Distribution. Southeast Asia: Cambodia, Laos, Thailand, Vietnam.

Senegalia delavayi (Franch.) Maslin, Seigler & Ebinger, Blumea 58: 40 (2013). - Acacia
delavayi Franch., PI. Delavay. 194 (1890).

Note. Two varieties recognised; both endemic to China.

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Card. Bull. Singapore 67(1) 2015

Senegalia delavayi (Fraiich.) Maslin, Seigler & Ebinger van delavayi - Acacia delavayi
Franch. van delavayi established by publication of A. delavayi van kunmingensis C.Chen &

H. Sun, Acta Bot. Yunnan. 12: 262 (1990).

Distribution. East Asia: China (Yunnan).

Senegalia delavayi van kunmingensis (C.Chen & H.Sun) Maslin, Seigler & Ebinger, Blumea
58: 40 (2013). - Acacia delavayi van kunmingensis C.Chen & H.Sun, Acta Bot. Yunnan. 12:
262 (1990).

Distribution. East Asia: China (Guizhou, Yunnan).

Senegalia donnaiensis (Gagnep.) Maslin, Seigler & Ebinger, Blumea 58: 40 (2013). - Acacia
donnaiensis Gagnep., Not. Syst. (Paris) 2: 114 (1911).

Distribution. Southeast Asia: Cambodia, Indonesia (Kalimantan), Malaysia (Sabah), Vietnam.

Senegalia ferntginea (DC.) Pedley, Bot. J. Linn. Soc. 92: 250 (1986). - Acacia ferruginea
DC., Prodr. 2: 458 (1825).

Distribution. Southeast Asia: Myanmar. South Asia: India, Sri Lanka.

Senegalia gageana (Craib) Maslin, Seigler & Ebinger, Blumea 58: 40 (2013). - Acacia
gageana Craib, Bull. Misc. Inform. Kew 1915: 409 (1915).

Distribution. Southeast Asia: Myanmar. South Asia: Bangladesh, Bhutan, India, Nepal,
Pakistan.

Senegalia intsia (L.) Maslin, Seigler & Ebinger, Blumea 58: 40 (2013). -Mimosa intsia L., Sp.
PI. 1: 522 (1753). -Acacia intsia (L.) Willd., Sp. PL, ed. 4, 4(2): 1091 (1806).

Distribution. Southeast Asia: Myanmar. South Asia: Bangladesh, India, Nepal, Sri Lanka.

Senegalia kekapur (I.C.Nielsen) Maslin, Seigler & Ebinger, Blumea 58: 40 (2013). - Acacia
kekapur I.C.Nielsen, Opera Bot. 81: 13, fig. 5 (1-8) (1985).

Distribution. Southeast Asia: Indonesia (Java, Sumatra), Singapore.

Note. Nielsen (1985a) noted that plants from NE Sumatra and Singapore may be taxonomically
different from S. kekapur sens. typ. from Java and southern Sumatra; this matter is currently
under investigation.

Senegalia kostermansii (I.C.Nielsen) Maslin, Seigler & Ebinger, Blumea 58: 40 (2013). -
Acacia kostermansii I.C.Nielsen, Opera Bot. 81:15 (1985).

Distribution. Southeast Asia: Indonesia (Alor Island, Sumbawa, Flores).

Senegalia meeboldii (Craib) Maslin, Seigler & Ebinger, Blumea 58: 40 (2013). - Acacia
meeboldii Craib, Bull. Misc. Infonn. 1928: 66 (1928).

Distribution. Southeast Asia: Myanmar, Thailand.

Senegalia megaladena (Desv.) Maslin, Seigler & Ebinger, Blumea 58: 41 (2013). - Acacia
megaladena Desv., J. Bot. Agric. 3: 69 (1814).

Note. Three varieties recognised.

Senegalia megaladena (Desv.) Maslin, Seigler & Ebinger var. megaladena - Acacia
megaladena Desv. var. megaladena established by publication of 4. megalodena var. garrettii

I. C. Nielsen and var. indo-chinensis I.C. Nielsen, Adansonia ser. 2, 19: 351 (1980).
Distribution. East Asia: China (Guangxi, Yunnan). Southeast Asia: Indonesia (Java), Laos,
Myanmar, Thailand, Vietnam. South Asia: Bangladesh, India, Nepal.

Synoptic overview oi Acacia sensu lato

245

Senegalia megaladena var. indochinensis (I.C.Nielsen) Maslin, Seigler & Ebinger, Blumea
58: 41 (2013). - Acacia megaladena var. indochinensis I.C.Nielsen, Adansonia ser. 2, 19: 351
(1980).

Distribution. Southeast Asia: Cambodia, Indonesia (Java, but probably introduced according
to Nielsen 1985a: 26), Laos, ?Malaysia (recorded for northern Peninsular Malaysia by Nielsen
1985a: 26 & 1992: 51, but not listed in Turner 1997: 290), Thailand, Vietnam.

Senegalia megaladena var. garrettii (I.C.Nielsen) Maslin, Seigler & Ebinger, Blumea 58: 41
(20X2)). - Acacia megaladena var. garrettii I.C.Nielsen, Adansonia sen 2, 19: 351 (1980).
Distribution. East Asia: China (Guangxi, Yunnan). Southeast Asia: Thailand.

Senegalia merrillii (I.C.Nielsen) Maslin, Seigler & Ebinger, Blumea 58: 41 (2013). - Acacia
merrillii I.C.Nielsen, Opera Bot. 81: 9, fig. 2 (1985).

Distribution. Southeast Asia: Indonesia (Moluccas, Sulawesi), Philippines.

Senegalia palawanensis (I.C.Nielsen) Maslin, Seigler & Ebinger, Blumea 58: 41 (2013). -
Acacia palawanensis I.C.Nielsen, Opera Bot. 81: 16, fig. 6 (7-11) (1985).

Distribution. Southeast Asia: Philippines.

Senegalia pennata (L.) Maslin, Nuytsia 22(6): 466 (2012). -Mimosa pennata L., Sp. PI. 1 : 522
(1753). -Acacia pennata (L.) Willd., Sp. PI, ed. 4, 4(2): 1090 (1806).

Note. Four subspecies recognised.

Senegalia pennata subsp. pennata. - Acacia pennata (L.) Willd. subsp. pennata established
by publication of A. pennata subspp. hainanensis (Hayata) I.C. Nielsen, insuavis (Lace)
I.C.Nielsen and kerri I.C.Nielsen, Adansonia ser. 2, 19(3): 352-353 (1980).

Distribution. Southeast Asia: Myanmar, Thailand (probably introduced). South Asia:
Bangladesh, Bhutan, India, Nepal, Sri Lanka.

Senegalia pennata subsp. hainanensis (Hayata) Maslin, Seigler & Ebinger, Blumea 58: 41
(2013). — Acacia hainanensis Hayata, Ic. PI. Fonuos. 3: 86 (1913). - Acacia pennata subsp.
hainanensis (Hayata) I.C. Nielsen, Adansonia ser. 2, 19(3): 352 (1980).

Distribution. East Asia: China (Fujian, Guangdong, Guangxi, Hainan, Yunnan). Southeast
Asia: My amnar, Vietnam. South Asia: India.

Senegalia pennata subsp. insuavis (Lace) Maslin, Seigler & Ebinger, Blumea 58: 41 (2013). -
Acacia insuavis Lace, Bull. Misc. Infonn. 1915: 401 (1915). -Acacia pennata subsp. insuavis
(Lace) I.C.Nielsen, Adansonia ser. 2, 19(3): 353 (1980).

Distribution. Southeast Asia: Cambodia, Laos, Myanmar. South Asia: India.
Introduced/cultivated. USA (Florida), Australia (Northern Territory, Queensland), Singapore,
Thailand.

Note. Pedley (2014) treated this entity as S. insuavis (Lace) Pedley; however, its taxonomic
status is currently under investigation by the present author.

Senegalia pennata subsp. ken ii (I.C.Nielsen) Maslin, Nuytsia 22(6): 467 (2012).

Acacia pennata subsp. kerrii I.C.Nielsen, Adansonia ser. 2, 19(3): 353 (1980).

Distribution. Australia: Queensland. East Asia: China (Yunnan). Southeast Asia: Cambodia,
East Timor, Indonesia (Java, Sulawesi, Flores, Komodo, Lombok, Sumbawa), Laos, Malaysia
(Peninsular Malaysia), Myanmar, Thailand, Vietnam. South Asia: Bhutan, India, Nepal, Sri
Lanka.

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Card. Bull. Singapore 67(1) 2015

Senegalia pluricapitata (Steud. ex Benth.) Maslin, Seigler & Ebinger, Blumea 58: 41 (2013).
Acacia pluricapitata Steud. ex Benth., London. J. Bot. 1:516 (1842).

Distribution. Southeast Asia: Indonesia (Sumatra, Java, Kalimantan), Malaysia (Peninsular
Malaysia, Sabah), ?Myanmar, Philippines, Thailand, Vietnam.

Senegalia pUiriglandulosa (Verde.) Maslin, Seigler & Ebinger, Blumea 58: 41 (2013). -
Acacia phiriglandulosa Verde., Kew Bull. 32: 472 (1978).

Distribution. Southeast Asia: Indonesia (West Papua), Papua New Guinea, Philippines.

Senegalia pruinescens (Kurz) Maslin, Seigler & Ebinger, Blumea 58: 41 (2013). - Acacia
pruinescens Kurz, J. Asiat. Soc Bengal, Pt. 2, Nat. Hist. 45: 296, 298 (1877).

Distribution. East Asia: China (Guangxi, Yunnan). Southeast Asia: Myanmar, Vietnam.

South Asia: India.

Senegalia pseudointsia (Miq.) Maslin, Seigler & Ebinger, Blumea 58: 41 (2013). - Acacia
pseudointsia Miq., FI. Ned. Ind. 1: 12 (1855), as ’"Pseudo-Intsia.

Distribution. Southeast Asia: Indonesia (Java, Sumatra), Malaysia (Peninsular Malaysia,
Sabah, Sarawak), Thailand.

Senegalia rugata (Lam.) Britton & Rose, N. Amer. FI. 23(2): 120 (1928). - Mimosa rugata
Lam., Encycl. 1: 20 (1783).

Acacia concinna (Willd.) DC., Prodr. 2: 464 (1 825). - Mimosa concinna Willd., Sp. PI., ed. 4,
4(2): 1039(1806).

Distribution. East Asia: China (Fujian, Guangdong, Guangxi, Guizhou, Hainan, Hunan,
Jiangxi, Yunnan). Southeast Asia: Cambodia, Indonesia (Ambon, Java, Kai Is., Sumba, Flores,
Moluccas, Sulawesi, Sumatra), Laos, Malaysia (Peninsular Malaysia), Myamnar, Papua
New Guinea, Philippines, Thailand, Vietnam. South Asia: Bangladesh, Bhutan, India (inch
Andaman Is.), Nepal.

Introduced. Australia (Queensland), Japan (Okinawa), Madagascar, Mauritius, Reunion Island.
Note. A variable species in need of critical revision.

Senegalia sulitii (I.C.Nielsen) Maslin, Seigler & Ebinger, Blumea 58: 42 (2013). - Acacia
sulitii I.C.Nielsen, Opera Bot. 81: 24, fig. 10 (6-10) (1985).

Distribution. Southeast Asia: Indonesia (Sulawesi), Philippines.

Senegalia tawitawiensis (I.C.Nielsen) Maslin, Seigler & Ebinger, Blumea 58: 42 (2013). -
Acacia tawitawiensis I.C.Nielsen, Opera Bot. 81: 22, fig. 10 (1-5) (1985).

Distribution. Southeast Asia: Philippines.

Senegalia teniana (Hanns) Maslin, Seigler & Ebmger, Blumea 58: 42 (2013). -Acacia teniana
Harms, Repert. Spec. Nov. Regni Veg. 17: 133 (1921),

Distribution. East Asia: China (Yunnan. Sichuan),

Senegalia thailandica (I.C.Nielsen) Maslin, Seigler & Ebinger, Blumea 58: 42 (2013). -
Acacia thailandica I.C.Nielsen, Adansonia ser. 2, 19: 356, pi. 1 (1980).

Distribution. Southeast Asia: Cambodia, Thailand.

Senegalia tonkinensis (I.C.Nielsen) Maslin, Seigler & Ebinger, Blumea 58: 42 (2013). -
Acacia tonkinensis I.C.Nielsen, Adansonia ser. 2, 19: 358, pi. 2 (1980).

Distribution. East Asia: China (Yunnan). Southeast Asia: Laos, Thailand, Vietnam.

Synoptic overview oi Acacia sensu lato

247

Senegalia torta (Roxb.) Maslin, Seigler & Ebinger, Blumea 58: 42 (2013). - Mimosa torta
Roxb., FI. Ind., ed. 2, 2: 566 (1832). -Acacia torta (Roxb.) Craib, Kew Bull. 1915: 410 (1915).
Distribution. Southeast Asia: ?Thailand. South Asia: India, Pakistan.

Note. It is unlikely that this species occurs in Southeast Asia. The Thailand records by Nielsen
(1985b: 167) are probably Senegalia tonkinensis (fide Srisanga & Sasirat, 2000).

Senegalia verheijenii (I.C.Nielsen) Maslin, Seigler & Ebinger, Blumea 58: 42 (2013). -Acacia
verheijenii I.C.Nielsen, Opera Bot. 81: 16, fig. 6 (1-6) (1985).

Distribution. Southeast Asia: hidonesia (Flores).

Senegalia vietnamensis (I.C.Nielsen) Maslin, Seigler & Ebinger, Blumea 58: 42 (2013). -
Acacia vietnamensis I.C.Nielsen, Adansonia ser. 2, 19: 360, pi. 3 (1980).

Distribution. Southeast Asia: Laos, Vietnam. (Erroneously recorded for China by Wu &
Nielsen 2010: 57).

Senegalia yunnanensis (Franch.) Maslin, Seigler & Ebinger, Blumea 58: 42 (2013). - Acacia
yunnanensis Franch., PI. Delavay. 193 (1890).

Distribution. East Asia: China (Yunnan, Sichuan).

VACHELLIA Wight & Am., Prodr. 272 (1834). TYPE: Vachellia farnesiana (L.) Wight &
Arn.

Delaportea Thorel ex Gagnep., Notul. Syst. (Paris) 2: 117 (1911). TYPE: Delaportea armata
Thorel ex Gagnep. (= Vachellia harmandiana (Pierre) Maslin, Seigler & Ebinger).

Nimiria Prain ex Craib, Bull. Misc. Infonn. 1927: 393 (1927). TYPE: Nimiria siamensis Craib
(= Vachellia siamensis (Craib) Maslin, Seigler & Ebinger).

Vachellia harmandiana (Pierre) Maslin, Seigler & Ebinger, Blumea 58: 42 (2013). -
Pithecolobium harmandianum Pierre, FI. Forest. Cochinch. 5: tab. 394A (1899). - Acacia
harmandiana (Pierre) Gagnep., Not. Syst. (Paris) 2: 115 (1911). - Delaportea armata Thorel
ex Gagnep., Not. Syst. (Paris) 2: 118 (1911).

Distribution. Southeast Asia: Cambodia, Laos, Thailand, Vietnam.

Vachellia inopinata (Prain) Maslin, Seigler & Ebinger, Blumea 58: 42 (2013). - Acacia
inopinata Prain, J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 66: 507 (1897). - Nimaria inopinata
(Prain) Craib, Bull. Misc. Infonn. Kew 1927: 393 (1927).

Distribution. Southeast Asia: Myamnar.

Vachellia kingii (Prain) Maslin, Seigler & Ebinger, Blumea 58: 42 (2013). - Acacia kingii
Prain, J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 66: 506 (1897).

Distribution. Southeast Asia: Myamnar.

Vachellia leiicophloea (Roxb.) Maslin, Seigler & Ebinger, Blumea 58: 42 (2013). - Mimosa
leucophloea Roxb., PI. Coromandel 2: 27, tab. 150 (1800). - Acacia leucophloea (Roxb.)
Willd., Sp. PI., ed. 4, 4(2): 1083 (1806).

Delaportea microphylla Gagnep., Bull. Soc. Bot. Fr. 99: 46 (1952).

Delaportea ferox Gagnep., Bull. Soc. Bot. Fr. 99: 47 (1952).

248

Card. Bull. Singapore 67(1) 2015

Distribution. Southeast Asia: East Timor, Indonesia (Java, Bali, Timor), Laos, Malaysia
(Peninsular Malaysia), Myanmar, Thailand, Vietnam. South Asia: India, Pakistan, Sri Lanka.
Introduced. Mauritius, Trinidad.

Note. Maslin et al. (20 1 3 : 42-43) recognise two varieties, VacheUia leucophloea var. leucophloea
and var. microcephcda (Kurz) Maslin, Seigler & Ebinger (recorded only for Myamnar). The
taxonomic status of VacheUia leucophloea var. microcephala, however, needs to be reviewed.

VacheUia myaingii (Lace) Maslin, Seigler & Ebinger, Blumea 58: 43 (2013). -Acacia myaingii
Lace, Bull. Misc, Infonn. 1915: 114 (1915), as “Myaingii”.

Distribution. Southeast Asia: Myanmar.

VacheUia nilotica (L.) P.J.H. Hurter & Mabb. subsp. indica (Benth.) Kyal. & Boatwr., Bot. J.
Linn. Soc. 172: 515 (2013). -Acacia arabica var. indica Benth., London J. Bot. 1: 500 (1842).
-Acacia nilotica subsp. indica (Benth.) Brenan, Kew Bull. 12: 84 (1957).

Distribution. Southeast Asia: Myanmar. South Asia: Bangladesh, India, Nepal, Pakistan.
West Asia: Iran, North Yemen, Oman, Qatar, South Yemen.

Introduced/naturalised. Africa, America, Australia (Northern Ten'itoiy, Queensland, South
Australia, Western Australia), India (Andaman Is,), Indonesia (Timor), Sri Lanka.

VacheUia siamensis (Craib) Maslin, Seigler & Ebinger, Blumea 58: 43 (2013). - Nimiria
siamensis Craib, Bull. Misc. Inform. 1927: 393 (1 927). -Acacia craibii I.C.Nielsen, Adansonia
ser.2, 19(3): 344 (1980).

Distribution. Southeast Asia: Thailand.

VacheUia tomentosa (Rottler) Maslin, Seigler & Ebinger, Blumea 58: 43 (2013). - Mimosa
tomentosa Rottler, Ges. Naturf. Freunde Berlin Neue Schriften 4: 208 (1803). - Acacia
tomentosa Willd. Sp. PL, ed. 4, 4(2): 1087 (1806).

Distribution. Southeast Asia: East Timor, Indonesia (Java, Sumba, Sumbawa, Sulawesi),
Myanmar, Thailand, Vietnam. South Asia: Bangladesh, India, Sri Lanka.

Appendix 2. Indigenous taxa of Acacia sensu lato in East and Southeast Asia, listed by
country of occurrence.

See Introduction for definition of the tenns East and Southeast Asia. E = endemic (additional
to the 1 5 countiy-specific endemics shown below there are 1 0 species of Senegalia that are
endemic to Southeast Asia which occur in more than one country, see text above).

East Asia

China

10 species
(13 taxa)

Senegalia caesia, S. catechu, S. delavayi (var. delavayi
& var. kunmingensis) [both E], S. megaladena (var.
megaladena & var. garrettii), S. pennata (subsp. hainanensis
& subsp. kerrii), S. pniinescens, S. rugata, S. teniana [E], S.
tonkinensis, S. yunnanensis [E].

Taiwan

2 species

Acacia confusa. Senegalia caesia.

Southeast Asia

Brunei

0

No indigenous species recorded.

Synoptic overview oi Acacia sensu lato

249

Cambodia

8 species
(9 taxa)

Senegalia caesia, S. comosa, S. donnaiensis, S. megaladena
van indochinensis, S. pennata (subsp. insuavis. & subsp.
kerrii), S. rugata, S. thailandica. Vachellia harmandiana.

East Timor

3 species

Senegalia pennata subsp. kerrii. Vachellia leucophloea, V.
tomentosa.

Indonesia

21 species
(22 taxa)

Acacia auriculiformis, A. mangium, A. oraria, A.
peregrinalis,A. wetarensis [E],4. sp. (Wetar) [E], Senegalia
borneensis, S. donnaiensis, S. kekapur, S. kostermansii
[E], S. megaladena (van megaladena & var. indochinensis
- probably introduced, see note above), 5*. merrillii, S.
pennata subsp. kerrii, S. pliiricapitata, S. pluriglandulosa,
S. pseudointsia, S. rugata, S. sulitii, S. verheijensii [E].
Vachellia leucophloea, V tomentosa.

Laos

9 species
(11 taxa)

Senegalia caesia, S. comosa, S. megaladena (var.
indochinensis & var. megaladena), S. pennata (subsp.
insuavis. & subsp. kerrii), S. rugata, S. tonkinensis, S.
vietnamensis. Vachellia harmandiana, V leucophloea.

Malaysia

8 species

Senegalia borneensis, S. donnaiensis, S. megaladena var.
indochinensis (uncertain, see note above), S. pennata subsp.
kerrii, S. pluricapitata, S. pseudointsia, S. rugata. Vachellia
leucophloea.

Myanmar

18 species
(21 taxa)

Senegalia caesia, S. catechu, S. chundra, S. ferruginea,
S. gageana, S. intsia, S. rneeboldii, S. megaladena var.
megaladena, S. pennata (subsp. hainanensis, subsp.
in.suavis, subsp. kerrii & subsp. pennata), S. pluricapitata
(uncertain), S. pruinescens, S. rugata. Vachellia inopinata
[E], V. kingii [E], V. leucophloea, V. myaingii [E], V. nilotica
subsp. indica, V. tomentosa.

Papua New
Guinea

10 species

Acacia auriculiformis, A. crassicarpa, A. leptocarpa.
Acacia mangium, A. peregrinalis, A. pubirhachis, A. simsii,
A. spirorbis subsp. solandri. Senegalia pluriglandulosa, S.
rugata.

Philippines

8 species

Acacia confusa. Senegalia merrillii, S. palawanensis [E], S.
pluricapitata, S. pluriglandulosa, S. rugata, S. sulitii [E], S.
tawitawiensis [E].

Singapore

1 species

Senegalia kekapur.

Thailand

17 species
(20 taxa)

Senegalia andamanica, S. caesia, S. catechu (uncertain), S.
comosa, S. rneeboldii, S. megaladena (var. indochinensis,
var. megaladena & var. garrettii), S. pennata (subsp. kerrii
& subsp. pennata), S. pluricapitata, S. pseudointsia, S.
rugata, S. thailandica, S. tonkinensis, S. torta (unlikely,
see note above). Vachellia harmandiana, V. leucophloea, V.
siamensis [E], V. tomentosa.

Vietnam

13 species
(15 taxa)

Senegalia caesia, S. comosa, S. donnaiensis, S. megaladena
(van indochinensis, van megaladena), S. pennata (subsp.
hainanensis and subsp. kerrii), S. pluricapitata, S.
pruinescens, S. rugata, S. tonkinensis, S. vietnamensis.
Vachellia harmandiana, V. leucophloea, V. tomentosa.

250

Card. Bull. Singapore 67(1) 2015

Table 1. Classification schemes for Acacia sensu lato showing species numbers and major
areas of occurrence.

Column 3 (in bold) gives the current classification and nomenclature of Acacia sensu lato.
Species numbers are sourced from the Worldwide Wattle website (web ref 4) plus 1 9 Australian
species of Acacia sensu stricto published in recent years but not yet posted to that site (see
Maslin, 2014a-d; Maslin & Bairett, 2014; Kodela, 2015); these numbers refer to accepted,
indigenous species only (not including infraspecific, infomial or hybrid formulae taxa).

Pre- Vienna IBC names
(Acacia treated as a single
genus with A. nilotica the
type)

Post- Vienna IBC names; Acacia sensu lato treated as five genera

With Acacia
nilotica the
type

With

Acacia
penninenns
the tj pe

Species numbers and distribution

Americas Africa*

Asia^

Australia

/Pacific

Total

Acacia

Acacia

VachelUa

57

843

30

9

1633

subgenus Acacia

subgenus Aculeiferum

section Spiciflorae

Senegalia

Senegalia

102

68^

45

2

201'*

section Filicinae

Acaciella

Acaciella

15

0

0

0

15

Acacia coidteri group

Mariosoiisa

Mariosousa

13

0

0

0

13

subgenus Phyllodineae

Racosperma

Acacia

0

03

13

1064®

1067

^Including Madagascar and Mascarenes. -New Guinea to the Middle East. ^Including three
species in Madagascar and one in Africa for which combinations are not yet available in
VachelUa. including seven species in Madagascar and two in Africa for which combinations
are not yet available in Senegalia. ^Acacia heterophylla was formerly recognised for Reunion
Island but was recently shown by Le Roux et al. (2014) to be conspecific with the Hawaiian
species A. koa and to have colonised the Mascarene archipelago dkectly from the Hawaiian
Islands < 1 .4 million yr ago. ^1 057 species in Australia, seven in the Pacific region.

Gardens’ Bulletin Singapore 67(1): 251-252. 2015
doi: 10.3850/S2382581215000198

251

BOOK REVIEW: Flowering Plants of the Western Ghats, Volume 1 (Dicots)
T,S, NayaVy A. Rasiya Beegam & M. Sibi. 2014 & Flowering Plants of the
Western Ghats, Volume 2 (Monocots). T.S. NayaVy M, Sibi & A, Rasiya
Beegam. 2014.

Thimvananthapuram: Jawaharlal Nehru Tropical Botanic Garden and Research Institute. 26 x
16.5 cm, hard cover, 1683 pp (Vols. 1. & 2). ISBN 978-81-920098-2-7 (Vol. 1) & ISBN 978-
81-920098-3-4 (Vol. 2). Price for the set of two volumes INR 3500 / US$ 200.

The mountain range of the Western Ghats,
whieh extends along the western coast of
the Indian subcontinent and exceeds 1600
km in length, is a well-known biodiversity
hotspot and a UNESCO World Heritage
Site. It is home to more than 8000 taxa
of flowering plants, of which about 15%
are endemic to the area. The Western
Ghats run through six states (from north
to south: Gujarat, Maharashtra, Goa,
Karnataka, Kerala and Tamil Nadu) and
although there are published checklists and/or Floras for each of these states, there
has not hitherto been a work focusing on the entire range. As such, this large and
comprehensive checklist of Flowering Plants is a very welcome addition to the
literature on Indian plants.

The checklist is published in two volumes. Volume 1 starts with a short
introduction to the Western Ghats, including basic information on the geography of
the region, rainfall, climate, major rivers, and forest types, together with a schematic
map. In the paragraph on methodology, the authors describe how they acquired and
verifled the information. The main body of Volume 1 is dedicated to an account of
Dicot families (arranged alphabetically) and genera (also arranged alphabetically, but
with cultivated taxa placed at the end of each family). Volume 2 deals with Monocots
in the same manner, but also includes two Addenda: additional taxa to be included
while the work was in production, and a list of species of doubtful occurrence. There
is also a list of references, and indices to scientiflc and local names. The index of local
names consists of over 13,000 entries and covers six languages spoken across the area:
Gujarati, Hindi, Kannada, Malayalam, Marathi and Tamil. The entire work is almost
1700 pages long.

The checklist is well-formatted, making full use of various fonts to provide
additional information (e.g. when the Western Ghats are highlighted in bold, it means
it is endemic to the area; and when the reference number is in italics it means that
the plant name is treated there as a synonym). The User’s Key explains well how the
entries are constructed and referenced, as well as explaining the formatting details,
making the work easy to use.

Each taxon entry starts with its accepted name, followed by any synonyms, its
habit and the availability of a description and illustration of the taxon. Information on

252

Card. Bull. Singapore 67(1) 2015

its complete native distribution is followed by its distribution and phenology within
the Western Ghats. These are followed by the threat category, economic importance
and a list of local names. Considering that this is just a checklist, it is surprisingly
well referenced. The list of references contains full citations to 2936 numbered works
with these numbers referenced in each entry to justify the information provided and to
allow the reader to follow up on details if needed.

A random check on families with which I am familiar shows that the authors
have strived to gather the latest literature to ensure that the names used are indeed, in
the vast majority of cases, up-to-date. However, I did spot some unusual placements
of genera, e.g. Cheilocostus and Costus of the family Costacaeae are placed in the
Zingiberaceae. The family Costaceae has been unambiguously recognised for at
least the last 50 years and has been reflected in all major recent works and databases,
including those used by the authors for verifications, so their family concept comes as
a bit of a surprise.

As a potential end user I regret that such a work as this, during an era of rapid
botanical exploration and publication, will inevitably become outdated in a few years,
and was not rather designed as an online project. Not only would that have allowed
various targeted searches to be performed, but the authors could also more dynamically
keep the data updated and respond to feedback. It may also perhaps have formed the
kernel of an expansion of the project into fully-fledged Flora of the Western Ghats,
which could also be illustrated. It would be lovely to see ferns and gymnosperms also
included.

Considering the extent of this work, the authors must no doubt be congratulated
on the final product, which must have required much perseverance and the ability to
cope with extremely large numbers of names, synonyms and references.

V

Jana Leong-Skornickova

Singapore Botanic Gardens